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MUSEUM OF VICTORIA

eagle

Records Of
The Museums & Art
Galleries of the
Northern Territory

Volume 12

December 1995

The BEAGLE, Records of the Museums and Art Galleries of the Northern Territory

(formerly “Records of the Northern Territory Museum of Arts and Sciences")

ABSTRACTED IN ZOOLOGICAL RECORD AND BIOSIS

Editorial Board

N. Smit Assistant Editor

U. Dasari Distributions

P.F. Murray

D. Megirian Editor

H.K. Larson Associate Editor

The Beagle is a refereed journal published by the Scientific Publications of the Museum and Art Gallery
of the Northern Territory to disseminate the results of research in the following areas:

(i) Systematic and other studies of the terrestrial, marine and freshwater flora and fauna of the
Northern Territory, tropical Australia, Southeast Asia and Oceania;

(ii) Australian Aboriginal, Southeast Asian and Oceanic arc material culture and archaeology;

(iii) Northern Territory and Indo-Paciftc history, maritime history and archaeology;

The Beagle is published once or twice a year, depending upon the material available and budgetary
considerations.

All contributions to The Beagle are peer reviewed by two referees, and where possible at least one
international referee is selected by the Editorial board.

Whilst articles for The Beagle will normally be 10-50 pages in length, shorter communications and
notes, book reviews and review articles may be acceptable also. Longer articles, significant works or
substantial revisions, which form integral studies, may be considered for separate publication in the
Monograph Series of the Museums and Art Galleries of the Northern Territory. Prospective authors
should contact the Editor, Scientific Publications.

The Beagle may be obtained by subscription or by exchange. Subscription rates for one year:
individuals $30.00, institutions $40.00. Postal charges, etc. In Australia only - $5.00; overseas - $10.00
(surface), $20.00 (airmail). Cheques should be made payable to ‘Northern Territory Museum’.

All subscriptions, back numbers and exchange enquiries should be addressed to the Librarian, Museum
and Art Gallery of the Northern Territory, PO Box 4646. Darwin, NT 0801, Australia.

AUTHORS OFFPRINTS

Twenty-five offprints are provided free for each published paper. Additional offprints may be
ordered when returning proofs.

SUBMISSION OF MANUSCRIPTS

Contributions, including the original and two copies of each manuscript, a computer floppy disk if
available, with copies of art work should be addressed to:

The Editor, Academic Publications

Museum and Art Gallery of the Northern Territory,

PO Box 4646, Darwin NT 0801,

AUSTRALIA.

ISSN 0811-3653

Government Printer of the Northern Terntory

CONTENTS

ERRATUM - Mislabelling of figures of holotypes in one of the Bruce papers in Vol. 11.ii

CLARK, L.G., WENDEL, J.F. AND CRAVEN, L.A. - A new species of Micraira (Poaceae:

Micraireae) from northern Western Australia.1

WILLAN, R.C. AND CATTANEO-VIETTL, R. - New data on Chelidoneura amoena Bergh,

1905 (Opisthobranchia: Cephalaspidea: Aglajidae).9

LARSON, H.K. - A review of the Australian endemic gobiid fish genus Chlamydogobius, with

description of five new species.19

WELLS, A. - Larva, pupa and notes on general biology of Tinodes radona Neboiss (Trichoptera:

Psychomyiidae).53

BRUCE, A.J. - Latreutes anoplonyx Kemp, 1914 (Crustacea: Decapoda: Hippolytidae), a jelly¬
fish associate new to the Australian fauna.61

LANSBURY, I. - Notes on the genus Anisops Spinola (Hemiptera - Heteroptera: Notonectidae)

of the Northern Territory and Western Australia.65

HORNER, P. - Two new species of Ctenolus (Reptilia: Scincidae) from the Northern Territory.77

AKERMAN, K. AND BINDON, P. - Dentate and related biface points from northern Australia..91

BRUCE, A.J. AND COOMBES, K. - The palaemonoid shrimp fauna (Crustacea: Decapoda:

Caridea) from the Cobourg Peninsula, Northern Territory.........101

MURANO, M. - Two new species of the genus Anisomysis (Crustacea: Mysidacea) from northern

Australia...145

BERGQUIST, P.R. AND KELLY-BORGES, M. - Systematics and biogeography of the genus

Ianthella (Demospongiae: Verongida: Ianthellidae) in the south-west Pacific.151

DEL CERRO, L. AND LLORIS, D. - A new species of Lepidotrigla (Scorpaeniformes:

Triglidae) from the waters off northern Australia.177

IZAWA, K. - A new fish parasite (Copepoda: Siphonostomatoida: Caligidae) from the Timor

Sea, Australia.185

MCINTOSH, I. - Who are the Bayinil .193

BOOK REVIEWS:

CARMENT, D. - Weary: the life of Sir Edward Dunlop.209

- "Red Ted": the life of E.G. Theodore.210

- Voyage from shame: the Cowra breakout and afterwards.210

- 25 April 1915: the inevitable tragedy.211

WELLS, J.T. - On the edge: women's experiences of Queensland.212

WILSON, H.J. - Creating a nation.213

Errata

BRUCE, A.J. 1994. Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific
alpheid shrimps of brevirostris species group. The Beagle, Records of the Museums and Art
Galleries of the Northern Territory 11: 15-28.

The caption to Figure 2 should read:

Fig. 2. Alpheus fenneri sp. nov. Allotype male, Sulawesi: A, first pereiopod, major chela, dorsal; B, same, minor
chela, dorsal. Juvenile male paratype, Sulawesi: C, scaphocerite; D, third maxilliped, lateral; E, first peri pod, major
chela, ventral; F, same minor chela, ventral.

The caption to Figure 5 should read:

Fig. 5. A (top left). Alpheus pubescens De Man, ovigerous female, Darwin Harbour, Northern Territory (photo.
A.J. Bruce); B (top right), Alpheus bellulus Miya and Miyake, female, Shirahama, Japan, (photo. Y. Miya);
C (bottom left), Alpheus williamsi sp. nov., ovigerous female holotype, off Charles Point, Northern Territory
(photo. A.J. Bruce); D (bottom right), Alpheus fenneri sp. nov., holotype, Menado, Sulawesi (photo. J.E. Randall).

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12 : 1-7

A NEW SPECIES OF MICRA1RA (POACEAE: MICRAIREAE) FROM
NORTHERN WESTERN AUSTRALIA.

LYNN G. CLARK 1 , JONATHAN F. WENDEL', AND LYN A. CRAVEN 2
1 Department of Botany, Iowa State University, Ames, Iowa 50011, USA.

2 Australian National Herbarium, Centre for Plant Biodiversity Research,

CS1RO Division of Plant Industry, G. P. O. Box 1600, Canberra, ACT 2601, Australia.

ABSTRACT

Micraira is an unusual genus of “resurrection” grasses endemic to Australia. During
fieldwork in northern Western Australia in 1993, vegetative material of a suspected
undescribed species of Micraira was collected. Examination of flowering material,
gathered from plants grown in cultivation, confirmed that this was indeed a new
species, which is here described and illustrated as Micraira lazaridis n.sp. A key to
the spicate species of Micraira is provided, and the new species is compared with the
other spicate taxa, using both gross morphological and leaf anatomical characters.

Keywords: Poaceae, Micraira, new species, identification key, Australia, Western
Australia.

INTRODUCTION

Micraira F. Muell. is a genus of typically
diminutive, moss-like, mat-forming grasses en¬
demic to Australia. The type species, M.
subulifolia F. Muell., is restricted to Queens¬
land, whereas the remaining 13 described spe¬
cies (including M. lazaridis) are narrow
endemics within the monsoon zones of the
Northern Territory and Western Australia
(Lazarides 1984). Species of Micraira are ex¬
traordinary in several respects. Perhaps most
notable is their ecological behavior as resurrec¬
tion plants, whereby they rapidly green and
flower from a desiccated state upon receiving
rainfall (Gaff and Latz 1978). In addition, the
genus exhibits spiral phyllotaxy and has an
unusual palea structure (Lazarides 1979,1984).
The most common habit adopted by the species
of Micraira is the mat form in which a dense
interlocking mat of stems is formed. On occa¬
sion, however, colonies on strongly fire-pro¬
tected pavements can develop into hummocks
up to c. 1 m tall.

During an expedition to Western Australia in
1993, additional vegetative material of a sus¬
pected undescribed species of Micraira was col¬
lected by J. Stewart, J. Wendel and E. Edwards.
Flowering material gathered from plants grown
in cultivation confirmed that this was indeed a
new species, which is here described and illus¬
trated. We deemed it appropriate to name this
species in honor of Michael (Mike) Lazarides,
who has studied this genus and described the
majority of its species. A key to the spicate
species of Micraira is provided, and the new
species is compared with other spicate taxa.

MATERIALS AND METHODS

Leaf anatomical studies were conducted in
addition to routine examinations and compari¬
sons of gross morphology. Leaf sections from
herbarium specimens were softened with Pohl’s
solution (Pohl 1965), hand-sectioned, and
mounted in lactophenol/aniline blue solution
(Sass 1958).

L. G. Clark, J. F. Wendel and L. A. Craven

SYSTEMATICS

Micraira F. Muell

Key to the spicate species of Micraira (adapted
from Lazarides 1984)

1. a Spike shorter than and obscured by the

upper leaves .. 2

b Spike prominently exserted above the up¬
per leaves. 3

2. a Leaf blades 5-6 mm long; glume apex

entire, scaberulous. M. inserta

b Leaf blades 1.8-3.7 mm long; glume apex
emarginate, ciliate. M. lazaridis sp. nov.

3. a Glumes terminating a flattened spine c.

0.5 mm long; leaf sheath abruptly nar¬
rowed at junction with bladeA/. compacta
b Glumes muticous; sheath of similar width
to blade at their junction.4

4. a Peduncle glabrous, glandular; florest c. 1/

2 as long as glumes; glumes oblong, obtuse,

glabrous, 1-3 nerved. M. spiciforma

b Peduncle pilose to hirsute, eglandular;
florest c. 1/3 as long as glumes; glumes

ovate, sub-acute, pilose, 1-nerved. M.

subspicata

Micraira lazaridis

L. G. Clark, Wendel & Craven, sp. nov.
(Figs 1-3, Plate 1)

Culmi aerii ramosi. Folia numerosa in
verticillis condensatis disposita; laminae 1.8-
3.7 mm longae, 0.3-0.4 mm latae, trinervatae.
Inflore-scentia spicata eglandulata foliis
superata. Spiculae 1-1.4 mm longae. Glumae
ovatae emarginatae ciliatae flosculis longiores.

TYPUS: Australia: Australian Capital Terri¬
tory: cu lti vated in greenhouse at Canberra, ACT,
7 September 1993, Craven 9344 (holotype:
CANB; isotypes: A, BRI, DNA, ISC, K, L,
MEL, NSW, PERTH). (Provenance: Western
Australia, sandstone pavement adjacent to cross¬
ing of King Edward River on road to Mitchell
Plateau. Voucher: Stewart, Wendel and Edwards
9253).

Other specimens examined. AUSTRALIA:
Western Australia: Mitchell Plateau, lat. 14°50'

5. long. 125°38'E, 26 February 1980, Dunlop
5312 (CANB); King Edward River at Mitchell
Plateau road crossing (Amax Crossing), lat. c.
14°53'S, long. 126°12'E, 2 June 1971, Byrnes
2196 (BRI, DNA), 29 February 1980, Dunlop

5363 (CANB), 1 March 1980, Dunlop 5415
(CANB), 26 May 1993, Stewart, Wendel and
Edwards 9253 (CANB, PERTH). Australian
Capital Territory: cultivated in greenhouse at
Canberra, 13 February 1987, Craven 8444
(CANB) (Provenance: Western Australia,
Mitchell Plateau road crossing of King Edward
River, leg. Craven, July 1984); ditto, 11 August
1993, Craven 9343 (CANB, ISC) (Provenance:
Western Australia, sandstone pavement adja¬
cent to crossing of King Edward River on road
to Mitchell Plateau, Stewart, Wendel and
Edwards 9253). UNITED STATES: Iowa: Story
Co.: Ames, campus of Iowa State University,
cultivated in R. W. Pohl Conservatory, 18 July
1992, Clark 1157 (ISC) (Provenance and voucher
details as for Craven 9343).

Description. Mat-forming perennial with
prop roots 3-8 cm long. Culms 0.5-0.9 mm wide
(with sheaths), 0.4-0.6 mm wide (without
sheaths), dark brown and smooth (below sheaths);
aerial culms 3-11 cm tall, extensively branched;
intemodes 1-1.5 mm long, completely covered
by the imbricate, persistent leaf sheaths. Leaves
numerous in tightly condensed, terminal whorls;
sheaths 0.8-1.0 (-1.5) mm long, as wide as blade
at their junction, 3-nerved, the midnerve well
developed and cartilaginous, the lateral nerves
thin, the intemerve areas green to hyaline, the
margins as broad as the strip bounded by the
nerves, membranous to hyaline, widening down¬
wards, enclosing the intemode at the base; blades
1.8-3.7 mm long, 0.3-0.4 mm wide, 2-3 times as
long as the sheath, subobtuse, 3-nerved, green
when young, adaxially flat, sparsely pilose be¬
tween the nerves, the midnerve as wide as the
margins, whitish, abaxially keeled, pilose be¬
tween the nerves, the nerves scabrous, the
midnerve prominent, whitish, twice as wide as
the margins, the margins cartilaginous, whitish,
scabrous; auricles absent; ligule a line of cilia ca.
0.2 mm long. Inflorescence a spike of sessile or
subsessile spikelets, 1.6-2.0 mm long, usually 5-
6 spikelets per spike, completely overtopped and
hidden by the whorl of leaves, subtended by a
single, keeled bract ca. 1.5 mm long; peduncle
eglandular, virtually absent; only pedicel of
terminal spikelet developed, this 0.2 mm long.
Spikelets 1-1.4 mm long, usually 2-flowered,
both florets apparently bisexual, laterally com¬
pressed; glumes 1-1.4 mm long, equal, longer
than the florets, membranous, ovate, obtuse,
emarginate, upper margins ciliate, l-or3-nerved,
the midnerve slightly wider than the somewhat

New resurrection grass

Plate. l.Micrairalazaridisn. sp. A. plant prior to commencement of irrigation; B, Plant seven
days after commencement of irrigation; C, Close-up of inflorescence, whorl is 7 mm in diameter.
(Material of Stewart et al. 9253 in cultivation. Photo credits: A, J. Faulkner; B, G. Brown; C,
S. Craig.)

L. G. Clark, J. F. Wendel and L. A. Craven

obscure lateral nerves and somewhat keeled, the
intemerve areas green to hyaline, the margins
broad, hyaline; lemmas 0.35-0.5 mm long, mem¬
branous; paleas 0.3-0.5 mm long, membranous.
Stamens 2; anthers 0.9-1.1 mm long, reddish-
purple before anthesis, becoming yellow at
anthesis, exserted above the leaf whorl. Stigmas
2, exserted before the anthers. Caryopsis 0.4
mm long, broadly ellipsoid.

Distribution and ecology. Western Australia:
the Mitchell Plateau area in the Kimberley re¬
gion (Fig. 1).

Recorded as occurring in shallow sand on
sandstone pavement. The flowering period in
nature is unknown.

Remarks. Micraira lazaridis is character¬
ized by its relatively tall, profusely branched
aerial culms; extremely condensed whorls of
leaves; three-nerved leaf blades 1.8-3.7 mm
long abaxially with a prominent, keeled
midnerve; highly reduced spicate inflorescence
not exserted above the upper leaves; and
emarginate.apically ciliate glumes that are longer
than the florets (Table 1). With respect to the
leaves and inflorescences, this species appears
to be among the most reduced within the genus,
although it has larger and more profusely

branched aerial culms than some of the other
species (Figs 2-3, Plate 1).

It is noted on the label for Craven 9343 that
•‘foliage became green two days after watering
commenced, and first flowers appeared five days
later.” Plants grown at Iowa State University
also greened and flowered from a desiccated
state within several days of initial watering. The
translucent stigmas exserted first, and withered
by the time the stamens shed their pollen. These
observations suggest that the species is
protogynous.

Renvoize (1986) examined transverse leaf
anatomy and leaf epidermal micromorphology
of two species, M. subulifolia and M. subspicata
Lazarides. He concluded that the leaf anatomy
sufficiently resembled that of the Arundineae to
warrant the inclusion of the Micraireae in the
same subfamily, the Arundinoideae. A starch
test confirmed the presence of the C, photosyn¬
thetic pathway. Weexamined the transverse leaf
anatomy of M. lazaridis and three other species
with spicate inflorescences for which material
was available. Renvoize (1986) noted that the
primary (or central) bundle was scarcely larger
than the other bundles, and this holds true in the
four species in our survey (Figure 3). He also

New resurrection grass

Fig. 2. Micraira lazaridis n.sp. A, habit; B, branch with leaf cluster and exserted stamens; C, leaf, adaxial view; D, leaf, abaxial
view; E, Inflorescence with subtending bract (arrow); F, spikelet with exserted stigmas and young stamens;
G.spikelet; H, glume, adaxial view. (A, based on material of Stewart el al. 9253 in cultivation; B-H, based on Clark 1157. Del.
J. Wojcik.)

L. G. Clark, J. F. Wendel and L. A. Craven

noted the presence of a broad sclerenchyma
girder below the bundles linking them to the
abaxial epidermis, and that girders above the
bundles could be absent, or, if present, could be
smaller than the lower girders. We observed this
in M. compacta Lazarides, M. spiciforma
Lazarides, and M. inserta Lazarides (Figure 3A,
C-D). In M. lazaridis , however, there are no
girders in direct contact with the bundles, al¬
though sclerenchyma is present (Figure 3B).

The four species shown here are easily distin¬
guished by their transverse leaf anatomy. Three
of them, M. lazaridis, M. inserta , and M.
spiciforma, have densely packed chlorenchyma
with no obvious intercellular spaces (Figure 3B-
D). The fourth species, M. compacta, shares the
densely packed chlorenchyma but has well de¬
veloped spaces on both sides of each bundle that
are reminiscent of fusoid cells in bamboos (Fig¬
ure 3A). The presence of a palisade-like ar¬
rangement in the chlorenchyma adjacent to the
adaxial epidermis was noted in M. compacta, M.
lazaridis, and M. inserta. Micraira lazaridis is
distinguished by its narrow blades with the
prominent keel, and the lack of any connection
between sclerenchyma bands and vascular bun¬
dles. Micraira inserta has a massive
sclerenchyma girder abaxial to the central bun¬
dle, but none of the other bundles have associ¬
ated girders; this species also apparently lacks

Fig. 3. Transverse leaf anatomy in four spicate species of
Micraira ; A. M. compacta (Craven 8337)', BM- lazaridis n.
sp .(Clark 1157); C, M inserta (Dunlop 4427)', D. M.
spiciforma (Dunlop 5298). Solid black - sclerenchyma.

obvious bulliform cells (Figure 3C). We suggest
that any future revision of Micraira include a
study of leaf anatomy, as this is likely to be
informative in circumscribing species and as¬
sessing relationships.

Without additional material and further study,
it is not possible to accurately infer relationships
among the species of Micraira. Based on simi¬
larities in leaf anatomy and the shared, reduced,
non-exserted spikes, however, M. lazaridis ap¬
pears to be closest to M. inserta. The phylogenetic
relationship of Micraira to other grasses is prob¬
lematical. Micraira is classified in its own tribe,
the Micraireae, which is placed in the
Arundinoideae (Clayton and Renvoize 1986;

Table 1. Comparison of the spicate species of Micraira.

Character

M. inserta

M. lazaridis

M. compacta

M. spiciforma

M. subspicata

Blade

persistence

deciduous

persistent

± persistent

persistent

Blade length
in mm

5-6

1.8-3.7

4-8

4-5

6-9

Sheath length
in mm

2-3

0.8-1.5

3-4

2-3

3-4

Ligule

ciliolate

0.3-0.5 mm

ciliolate
c. 0.2 mm

tubercular-
thickened ridge,
glabrous or sparsely
puberulous

membranous or
tubercular, pube¬
rulous or
glabrous

Inflorescence
length in mm

3-4

1.6-2

4-5

3-4

Inflorescence
exsertion at
maturity

not exserted

well exserted

shortly

exserted

Presence of
glands in spike

eglandular

glandular

eglandular

Glume apex

entire,

scaberulous

emarginate,

ciliate

emarginate,

scaberulous

entire or minutely
emarginate, muticous,
minutely scaberulous

entire,

muticous

New resurrection grass

Watson and Dallwitz 1992) or segregated into
its own subfamily, the Micrairoideae (Lazarides
1979). Clayton and Rcnvoize (1986, p.184)
speculate that Micraira is a “derivative of
Arundineae adapted to a specialized habitat.” In
this respect, we note that in a cladistic analysis
of the grass family based on DN A sequence data
from the chloroplast gene ndhF, M. lazaridis
consistently appears as sister to a robust clade
containing the four representatives of the
Chloridoideae included in the survey (Clark et
al. in prep.). Further study is required, but it is
likely that Micraira is not closely related to the
Arundineae.

ACKNOWLEDGMENTS

The authors gratefully acknowledge the fi¬
nancial support of the National Geographic So¬
ciety (to JFW) and the National Science Founda¬
tion (to LGC and J FW). The co-operation of Mac
Stewart and Ted Edwards in the collection of
specimens is appreciated.

REFERENCES

Clayton, W. D. and Renvoize, S. 1986. Genera
graminum. Her Majesty’s Stationery Office: Lon¬
don.

Gaff, D. F. and Latz, P. K. 1978. The occurrence of
resurrection plants in the Australian flora. Aus¬
tralian Journal of Botany 26: 485-492.

Lazarides, M. 1979. Micraira F. Muell. (Poaceae,
Micrairoideae). Brunonia 2: 67-84.

Lazarides, M. 1984. New taxa of tropical Australian
grasses (Poaceae). Nuytsia 5(2): 273-303.

Pohl, R. W. 1965. Dissecting equipment and materi¬
als for the study of minute plant structures.
Rhodora 67: 95-96.

Renvoize, S. 1986. A survey of leaf-blade anatomy in
grasses VIH. Arundinoideae. Kew Bulletin 41:
323-338.

Sass, J. E. 1958. Botanical microtechnique. Iowa
State University Press: Ames, Iowa.

Waston, L. and Dallwitz, M. J. 1992. The grass
genera of the world. CAB International:
Wallingford, Oxon, England.

Accepted 22 July, 1995

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:9-18

NEW DATA ON CHELIDONURA AMOENA BERGH, 1905
(OPISTHOBRANCHIA: CEPHALASPIDEA: AGLAJIDAE).

R. C. WILLAN 1 AND R. CATTANEO-VIETTP
1 Museum and Art Gallery of the Northern Territory,

GPO Box 4646, Darwin, NT 0801, Australia.

2 Istituto di Zoologia, Universita di Genova,
via Balbi 5, 16126 Genova, Italy.

ABSTRACT

Chelidonura amoena Bergh, 1905 is redescribed, principally on material from
Australia. The single extant type specimen is designated as lectotype. Considerable
intraspecific colour variation is evident, with that relating to the caudal lobes being
greatest. The variation in coloration and shape observed in one specimen from Belau
lies outside those described for C. amoena and we believe it could be explained as the
product of hybridisation with C. electro Rudman, 1970. Aggregation for the purpose
of breeding is shown to be normal for C. amoena. The collection data and observations
indicate C. amoena is widely distributed in the tropical western Pacific and eastern
Indian Oceans with disparate limits on the eastern and western coasts of the Australian
continent.

Keywords: Mollusca, Opisthobranchia, Cephalaspidea, Aglajidae, Chelidonura, tax¬
onomy, intraspecific variation, hybridisation.

INTRODUCTION

Chelidonura amoena Bergh, 1905 is a little-
known member of the Aglajidae, a family of
advanced, carnivorous, cephalaspidean
opisthobranchs. The species was described from
two specimens collected in 1899 at “Saleyer” (=
Selayar) Island, southern Sulawesi, Indonesia,
during the “Siboga” Expedition and it was not
reported again until Coleman (1975) illustrated
it in his popular book What Shell is That ?. That
report was based on a specimen from Coral Bay,
central Western Australia, found in 1972. Ho
(1989) wrote an illustrated account about a speci¬
men from Pulau Kapas, Malaysia. More recently,
photographs of specimens from Western Aus¬
tralia have appeared in books by Wells et al.
(1990) and Wei Is and Bryce (1993), and the name
has been used in taxonomic (Rudman 1970,1974;
Gosliner 1980) and faunal (Wells and Slack-
Smith 1986) checklists. However,at no time have
authors actually compared their material with
the original specimens, or has the species been
redescribed on live or freshly collected material.

Spurred by finding one specimen on the Great
Barrier Reef, Queensland, Australia, we have
accumulated data on external morphology, in¬
ternal anatomy (shell, buccal bulb, penial com¬
plex), reproductive behaviour and geographic
range pertaining to this distinctive and inter¬
esting opisthobranch. The extent of intraspecific
variation in coloration in this species has turned
out to be considerable - seemingly greater than
that recorded in any other species of Cheli¬
donura. One animal from Belau (= Palau) pos¬
sibly represents a hybrid with C. electro
Rudman, 1970. Finally, the geographic range
of C. amoena has turned out to be much more
extensive than previously appreciated, and the
species has proven to be relatively common in
parts of this range.

Abbreviations used in the text are as follows:
AMS, Australian Museum, Sydney; NMV, Mu¬
seum of Victoria, Melbourne; NTM, Museum
and Art Gallery of the Northern Territory, Dar¬
win; WAM, Western Australian Museum, Perth;
ZMA, Zoologisch Museum, Universiteit van
Amsterdam, the Netherlands.

R.C. Willan and R. Cattaneo-Vietti

SYSTEMATICS

Family Aglajidae Pilsbry, 1895
Genus Chelidonura A. Adams, 1850
Chelidonura amoena Bergh, 1905
(Plate 1, Figs 1-4)

Chelidonura amoena Bergh, 1905:45,46, pi. 3,
fig. 7. - Rudman 1970:10; - Rudman 1974:207; -
Coleman 1975:107,no.312;-Gosliner 1980:333;
- Wells and Slack-Smith 1986:52; - Ho 1989:58,
59; - Wells et al. 1990: 66, pi. 43, no. 303; Wells
and Bryce 1993: 36,37, species number 28.

Through the courtesy of Mr R. Moolenbeek,
ZMA, we have been able to examine the type
material of Chelidonura amoena Bergh. (All the
existing type material of species described by
Bergh from the Siboga Expedition is deposited
in ZMA.) The type material consists of an intact,
18 mm specimen, plus a 6 mm long piece of
black, sickle-shaped tissue. The intact animal is
unquestionably the smaller of Bergh's syntypes,
and we herein designate it as lectotype. Appar¬
ently Bergh completely disposed of the larger
syntype after dissecting it because the piece of
black tissue cannot be matched with any exter¬
nal structure or internal organ mentioned in the
original description; it is probably only a con¬
taminant.

Type material. LECTOTYPE. Here desig¬
nated. Intact preserved specimen (18 mm long
by 5 mm maximum width), with tail bent to the
left side, collected on (presumably low-tidal)
reef, “Saleyer” (= Selayar) Island, southern
Sulawesi, Indonesia - 06°00’S, 120°30’E, 26
October 1899 (Siboga Expedition, Station 213).
Deposited in ZMA.

Additional material. QUEENSLAND - One
specimen (Fig. 1), 30 mm extended crawling
length, 10 metres depth, Davies Reef, N. E. of
Townsville, central section. Great Barrier Reef,
Queensland, Australia - 18°50’S, 147°39’E, R.
Cattaneo-Vietti, 13 November 1986 (Italian
Great Barrier Reef Expedition, Station 30.) Speci¬
men observed and photographed live by R.
Cattaneo-Vietti, dissected by R. C. Willan. Dis¬
sected body in AMS C166226. WESTERN
AUSTRALIA -Two specimens,crawling lengths
not recorded, depths not recorded, western side
of North Lagoon, Scott Reef, northern Western
Australia - 14°00’S, 121°45’E, C. W. Biyce,
September 1984, WAM 2417-84; two speci¬
mens, 21, 16 mm crawling length, 1.2 metres

depth, Bernier Island, Dampier Archipelago,
northern Western Australia-24°52’S, 113°08’E,
C. W. Bryce, March 1980, WAM 2417-84; three
specimens, crawling lengths of the larger two
recorded as 26 and 21 mm, photographed in situ,
subtidally, Steep Point, Shark Bay, central West¬
ern Australia - 26°08’S, 113° 10’E, C.W. Bryce,
March 1986, WAM 345-85; one specimen, 35
mm crawling length, on crustose algae, 2 metres
depth. Coral Bay, south of Learmonth, central
Western Australia - 23°00’S, 113°49’E, N.
Coleman, 14 July 1972, NMV. (Mr Bryce kindly
made available a list of 105 specimens of this
species in WAM. These specimens were col¬
lected from Ashmore Reef, Timor Sea, south to
Rottnest Island, Western Australia.between 1980
and 1986.)

Photographic material. WESTERN AUS¬
TRALIA - Many individuals (some shown here
in Plate 1), crawling lengths not recorded, 6
metres depth, western side of Mamey Bay, Malus
Islands, Dampier Archipelago, northern West¬
ern Australia - 20°31’S, 116°4rE, W. R.
Rowlands and G. McGarvie, 4 November 1990,
four colour transparencies examined; one speci¬
men, 16 mm crawling length, photographed in
situ, subtidally, Rottnest Island, southern West¬
ern Australia - 32°00’S, 115°30-E, C. W. Bryce,
November 1980, colour transparency examined.
One specimen, 24 mm crawling length, photo¬
graphed in situ, subtidally, Rottnest Island, south¬
ern Western Australia - 32°00'S, 115°30’E, C.
W. Bryce, December 1981, colour transparency
examined. BELAU (= PALAU) - Two speci¬
mens (Fig. 4), crawling lengths not recorded.

The opisthobranch Chelidonura amoena in Australia

selected from a breeding population in a sandy
area on the edge of Rengrak Channel, Belau -
7°30’N, 134°30’E, C. Carlson, 28 June 1969,
colour transparency examined. RYUKYU IS¬
LANDS - Two specimens, both approximately
30 mm extended crawling length, collected
intertidally on Oujima Island, Okinawa -
26°21’N, 127°36’E, I. Hamatani, 4 November
1991, colour print of one specimen examined.

Description. (Based on Queensland speci¬
men.) Living animals illustrated in Plate 1.

Extended length to 40 mm (Ho 1989), though
30-35 mm is more usual for adults.

When alive, body shape corresponded to
Rudman’s (1974: 201-202) generic diagnosis.
When crawling actively, parapodia occupied
60% of entire body length (measured from ante¬
rior margin of head shield to posterior tip of left
caudal lobe).

Anterior margin of head shield broad, trun¬
cate, bluntly trilobed with no mounds or sensoiy
bristles visible. Head shield tapering gradually
toward pointed posteriorend beneath parapodia.
Anterior margin of foot slightly enlarged, its
margins folding outwards, but never forming
funnels. Parapodia relatively low, touching one
another only in dorsal midline and not overlap¬
ping. Both caudal lobes elongate, flattened left
lobe almost three times length ofdorsally keeled
right lobe.

Head shield pale grey on account of numer¬
ous, microscopic, evenly spaced black specks,
darkest mid-dorsally. Interspersed with these
black specks are numerous, fine, white flecks.
Anterior margin ofhead shield, including fronto-
lateral comers, marked with broad, pale cream
band. Foot sole uniformly grey. Parapodia ap¬
pearing uniformly grey-black to naked eye, but
magnification reveals fine, black reticulum, dark¬
est closest to midline. Black pigmentation shad¬
ing to pale cream-gold margin that is narrower
than cream band on front of head shield. This
pale margin separated from black area by a very
irregularline. Posteriorshield dark grey dorsally,
equivalent in intensity to head shield, speckled
with tiny pallid areas but not reticulate. Dark
grey pigmentation of mantle shield continuing
without interruption onto dorsal surfaces of both
caudal lobes. Margins of caudal lobes cream-
gold, sharply delineated from central dark area
of mantle shield. Apices of caudal lobes with no
yellow pigmentation.

When examined after preservation in ethanol,
body pale grey, darkening on antero-lateral cor-

Fig. 2. Chelidonura amoena shell from specimen shown in
Figure I, length 4.9 mm. A, exterior, B, interior.

ners and posterior margin of head shield,
parapodial margins, and dorsal ridges of caudal
lobes. Interior of parapodia white.

Shell (Fig. 2) with dimensions 4.9 x 2.9 mm,
located internally underneath posterior shield,
calcareous but thin, white, consisting of calci¬
fied apex and greatly expanded, flattened body
whorl. Protoconch and spire whorls heavily
calcified, with superficial microsculpture of ir¬
regular, small pustules and ridges. Body whorl

R.C. Willan and R. Cattaneo-Vietti

relatively large, subrectangular, calcified, exte¬
rior glossy from membranous, overlying perio-
stracum, interior shallow and dull white. Ante¬
rior margin broadly rounded and lightly calci¬
fied. Posterior margin short, deeply concave,
defined by calcified ridge internally, produced
into long, erect, narrow spine, elevated above
level of protoconch. Shell’s right (i.e. columellar)
margin evenly convex. Shell surface ornamented
externally with approximately 20, strong, broad,
flattened concentric ridges. Ridges all of vari¬
able width, all becoming narrower toward pos¬
terior margin.

Buccal bulb3 mm in length (i.e. approximately
14 preserved body length excluding caudal lobes),
forming most conspicuous organ within ante¬
rior visceral cavity, barrel-shaped, elongate, par¬
allel-sided, consisting of three, large, longitudi¬
nal blocks of muscle - two lateral, and one, a
little shorter, mid-ventral. Inner walls of all 3
muscle blocks completely smooth. Salivary
glandsshort, located at extremehind end of buccal
bulb, ducts opening either side ofventral muscle
block. No vertical sheet of tissue (diaphragm)
separating crop and organs of visceral mass.

Penial complex (Fig. 3). Penis eversible, lo¬
cated inside head cavity to right of mouth and
below right fronto-lateral comer of head shield.
Penial complex unpigmented, narrower than
and only 2/3 length of buccal bulb beside which
it lies. When retracted, penial complex elongate
and circular, consisting of two sections of about
equivalent length. Proximal section, the penial
sheath (= vestibule) containing penial papilla,
fusiform and thin-walled. Upon entering this
sheath, ventral fold of external seminal groove
enlarges threefold, forming inner wall of deep

interior seminal groove located on floor to right
of papilla; this internal groove runing directly to
base of papilla. Penial papilla Vi length of sheath,
muscular and conical, flattened, margins en¬
folded to form deep ventral groove for entire
length. Distal part of penial complex (i.e.
prostatic section) fusiform and circular, nar¬
rower and slightly longer than sheath, tapering
very gradually to bluntly rounded apex where
two retractor muscles insert dorsally. Walls of
prostatic section thick and glandular, but not
distinctly granular. Tall ridge passing through
interior of prostatic section. Depression on right
of ridge continuous with both internal seminal
groove and groove on outside of penial papilla.
Penial retractor muscles short and broad, origi¬
nating from floor of visceral cavity.

DISCUSSION

Colour variation. The specimens and photo¬
graphs of Chelidonura amoena now available
indicate that the species possesses considerable
colour variation. The head shield and posterior
shield are relatively constant in colour, whereas
the parapodia and caudal lobes are variable from
individual to individual.

With the exception of one of the animals
from Palau (see below), the head shield is grey¬
ish black with white speckles, becoming paler
anteriorly, and merging gradually into the rela¬
tively broad cream-gold to yellow-gold band
that borders the front of the head. This differs
from Bergh’s (1905) original illustration of a
pale, longitudinal, medial bar extending rear¬
ward from this band into the dark area. No

penial papilla

prostate gland

cut wall

of penial sheath

penial retractor
muscles

Internal seminal
groove

external body
wall

0.5 mm

Fig. 3. Chelidonura amoena penial complex of specimen shown in Figure 1. Sheath cut away dorsally to reveal internal seminal
groove and papilla.

The opisthobranch Chelidonura amoena in Australia

animal resembling Bergh's illustration has ever
been seen again and the illustration is prob¬
ably inaccurate. Further, the golden margin
along the front of the head would probably
have been more vivid than the depiction would
indicate.

The parapodia are dark dorsally with quite
vivid yellow-gold anterior comers. The parapodia
are generally black and peppered with white to
the extreme margin (Plate 1). In the Queensland
specimen, a narrow, white line clearly demar¬
cates the parapodial margins (Fig. 1). We have
never seen a spec imen match i ng that depicted by
Bergh with a relatively broad white margin to
the parapodia. The extent of black pigment on
the parapodia is quite variable. Generally, they
become lighter gradually, through a reticulate
intervening section, to the pale grey foot, but in
the specimen depicted by Bergh and in WAM
2417-84, only the upper half of the parapodia is
black. Actually, in WAM 2417-84, the upper
half of the right parapodium is black and there
are only two large blotches on the left parapodium
(see Wells and Bryce 1993: 36).

The posterior shield is usually uniformly dark
(pale grey to sooty black) with microscopic
white speckles. In approximately half the ani¬
mals, there is an unpigmentedareaon either side
resulting in an hour-glass pattern lying across
the top of the shield. This marking is well shown
in Bergh’s animal. The specimen illustrated by
Coleman (1975) is unique in having an opaque
white shield with a single, large, black blotch
mid-dorsally.

The pigmentation of the caudal lobes is the
most variable element of this species’ colora¬

tion. In the Queensland specimen, the caudal
lobes are dark throughout their length (both
inside and outside), with a narrow, dull cream
margin. In the specimens illustrated by Bergh
(1905), Ho(1989) and Wells et al. (1990), and
in the smaller animal photographed in Palau
(Fig. 4A), these lobes are black anteriorly, pale
cream marginally and vivid gold apically. The
caudal lobes of Coleman’s specimen are opaque
white with cream-gold tips. Those of the
Okinawan specimen are cream with a narrow,
black stripe on the dorsal face of the left lobe.

The larger animal from Belau (Fig. 4B) seems
to lie outside the range of colour variation just
described for Chelidonura amoena in several
ways. Its entire head shield was translucent
white. Its parapodia were translucent white with
yellow margins and, symmetrically, irregular
black blotches on the anterior third. Its posterior
shield was opaque white. Both its caudal lobes
were opaque white with pale yellow margins.
This specimen was part of a breeding aggrega¬
tion of C. amoena (see below), and it was photo¬
graphed deliberately because its pale coloration
rendered it conspicuous amongst the mass of
otherwise dark individuals (C. Carlson pers.
comm. 1990). Unfortunately the specimen was
not retained by the collector.

Several explanations could account for this
anomalous individual. It could have been an
extreme colour form of Chelidonura amoena ,
but in view of the hundreds of specimens of that
species now known that seems unlikely, as the
individual lies too far outside the variation of
that species. Secondly, it could have been a
specimen of the closely related C. electra Rudman

Fig. 4. Chelidonura amoena. Specimens from breeding aggregation at Rengrak Giannel, Belau, 28 June 1969, lengths not
recorded. Drawn to same scale from slide by C. Carlson. Specimen 4B is possible C. amoena x C. electra hybrid. Black
pigmentation represented by heavy stipple; gold or yellow marginal lines represented by lighter stipple.

R.C. Willan and R. Caitaneo-Vietti

Plate 1. A, Chelidonura amoena. Actively crawling specimen separated from the breeding aggregation at Dampier Archipelago
for photography, 4 November 1990, length not recorded. Reproduced from slide by G. McGarvie.

B, specimens from the breeding aggregation at Dampier Archipelago, 4 November 1990. Note the small individual to the left
of centre. Reproduced from slide by G. McGarvie.

The opisthobranch Chelidonura amoena in Australia

1970, that had been attracted by pheromones
released from all the C. amoena in the aggrega¬
tion. The characters of C. electro exhibited by
this specimen (in contrast to C. amoena) are its
longer body, longer and narrower head shield,
translucent areas, pale yellow (not gold) bor¬
ders, opaque white posterior shield and caudal
lobes, and longer, taller, and more attenuated
caudal lobes. Chelidonura electro has not yet
been recorded from Belau, but little intensive
collecting has been done there to this date (C.
Carlson pers. comm. 1992). Thirdly, and in our
opinion the most likely scerario, is that this
individual was hybrid between C. amoena and
C. electro because it possesses pigmentation and
shape characteristic of both species.

Hybridisation has already been strongly sus¬
pected in prosobranch gastropods (e.g.
Springsteen and Leobrera 1986: 69; Marshall
1995: 85,87), but it is not presently recognised
as occurring in any opisthobranch.

We conclude that colour variation is consider¬
able in Chelidonura amoena and this variation
can neither be correlated with geographical oc¬
currence nor interpreted as clinal. For example,
two of the Australian specimens (the one illus¬
trated by Coleman (1975) and the specimen from
Queensland) differ more from each other than
either does from the Indonesian one depicted by
Bergh (1905). The largest population for which
data on colour variation is available is that from
the Dampier Archipelago where an aggregation
of dozens of individuals was encountered in
November 1990. Members of this population
displayed most of the colour variation described
above for the species as a whole (W. R. Rowlands
pers. comm. 1994). Data on large populations
such as this are unavailable at present to quan¬
tify the components of chromatic variation.

Comparison with similar species. The ge¬
nus Chelidonura is circumtropical and consists
of approximately 20 species (Rudman 1973,
Perrone 1990, Martinez et al. 1993, Yonow
1994). R.C.W. has had the opportunity to study
living specimens of seven other Indo-Pacific
species besides C. amoena: C. electro ; C.
fulvipunctata Baba, 1938 (= C. conformata Bum,
1966; for synonymy, see Gosliner 1980: 334,
335); C. hirundinina (QuoyandGaimard, 1832)
(= C. philinopsis El iot, 1903; for synonymy, see
Gosliner 1987: 42 and Gosliner 1988: 91); C.
inornata Baba, 1949; C. pallida (Risbec, 1951);
C. tsurugensis Baba and Abe, 1959; and C.
varians Eliot, 1903.

In comparison to Chelidonura amoena, C.
electro is larger when adult (to 52 mm extended
crawling length), more elongate, more translu¬
cent, and there is a pale yellow (not gold) edging
to the head shield, parapodia and caudal lobes.
Dark forms of C. amoena, like the Queensland
specimen and the one illustrated by Wells et al.
(1990), resemble C. inornata and C. tsurugensis.
However, C. inornata has a uniform soot-black
head shield with a broad, opaque white frontal
band and orange antero-lateral comers, soot-
black parapodia with white spots and a narrow
white band, and uniform soot-black posterior
shield and caudal lobes. Chelidonura tsurugensis,
although exceedingly variable in coloration it¬
self, is most often uniformly black. Chelidonura
tsurugensis can always be distinguished from all
the other Chelidonura species by its reduced
right caudal lobe.

R.C.W. had previously considered partition¬
ing the species of Chelidonura into two species
groups. One character contemplated for this
separation was pigmentation. It seemed possible
to recognise a group with translucent, colourless
bodies (e.g. C. pallida and C. electro) and a
second group with heavily pigmented, black or
dark brown bodies (e.g. C. inornata, C.
fulvipunctata and C. tsurugensis). However, in
possessing great variability in pigmentation on
the parapodia and caudal lobes, C. amoena
could be placed in either group depending on the
intensity of its pigmentation. Such an observa¬
tion forces us to reject the hypothesis.

The wording by Risbec (1928:42), translated
from the French: “Entire body and parapodia
displaying a conspicuous greenish-maroon net¬
work over a yellow background” initially led us
to consider the possibility that Doridium
perparvum Risbec, one of the most enigmatic
of aglajid names (Rudman 1974: 209; Gosliner
1980: 284), might have been a junior synonym
of Chelidonura amoena based on a dark colour
form. However, other characters contained in
Risbec’s (1928) original description and subse¬
quent revised account (Risbec 1951: 134, pi. 8,
fig. 8; as Chelidonuraperparava [sic]) force us
to discount this possibility. The shell of D.
perparvum is neither calcified nor does it have
a spine on its posterior margin. The buccal bulb
of D. perparvum is “enormous” and spherical,
and its lining possesses numerous, glistening,
soft hooks (Risbec (1928) used the term “papil¬
lae” to describe them in the legend to his Fig¬
ure 18). None of these characters is concordant

R.C. Willan and R. Cattaneo-Vietti

with C. amoena and there is no justification for
synonymising D. perparvum with this species.
Indeed, D. perparvum may not even be a spe¬
cies of Chelidonura, because according to
Gosliner (1980: 334), the shell of all species in
that genus should be “thinly but completely
calcified”. The identity of D. perparvum re¬
mains unresolved.

Breeding aggregations. We are aware of
three instances of breeding aggregations in
Chelidonura amoena, only one of which has
been mentioned previously in the literature. On
each occasion, observers discovered aggregations
of many individuals, all forming clusters. Within
these clusters, individuals were crawling over
each other and/or laying spawn. In June 1969,
Mr C. Carlson observed such an aggregation
intertidally at Rengrak Channel, Belau. He came
across 50 to 70 animals in a sandy area on the
edge of the channel. The presence of spawn was
interpreted as indicating a breeding aggrega¬
tion. Wells and Bryce (1993: 37) reported that
mating and spawning individuals were found at
South Passage, Shark Bay, in March 1986. In
November 1990, Messrs W. Rowlands and G.
McGarvie observed an aggregation subtidally at
Malus Islands, Dampier Archipelago. All the
specimens in this aggregation were crawling
actively in tight clusters of up to a dozen indi¬
viduals (Plate lb). The only growth on the silty
rocks on which the specimens were amassed was
a low stubble of filamentous algae. Some ani¬
mals were actually laying masses of spawn when
the aggregation was observed, so this behaviour
was interpreted as a mass spawning event. The
spawn consisted of a white tangle inside a clear,
flaccid sac. Interestingly, there were a few very
small animals less than half the size of the
majority amongst the clusters (Plate 1). Either
these animals were diminutive adults that were
engaged in the spawning, or else they were non-
reproducing juveniles attracted by the
pheromones emitted by all the other mature
animals.

Since discrete mating pairs have not been
observed in Chelidonura amoena, we assume
that aggregation for breeding always takes place
in this species. Aggregation as a normal part of
the reproductive process has already been re¬
corded for Philinopsis cyanea (Martens), and
one other member of the Aglajidae (Allan 1950:
217). This phenomenon of aggregation for breed¬
ing has also been well documented in sea hares

(Allan 1950: 21 l,Kandel 1979,Carefoot 1987).

Spawning events could take place more than
once in a year in Chelidonura amoena because
the two sets of observations on the coast of north¬
ern Western Australia were five months apart.

Geographical distribution. Far from being
restricted to Indonesia, as the original record of
Chelidonura amoena might indicate, it is now
apparent that this species occurs widely through¬
out the western Pacific (i.e. Ryukyu Islands,
Belau, eastern Australia) and eastern Indian
Oceans (i.e. Western Australia, Christmas Is¬
land, Indonesia, Malaysia). The northern bound¬
ary apparently lies south of Japan and Guam
because no specimens have been found at either
locality despite many years of intensive
opisthobranch study (K. Baba pers. comm. 1990,
C. Carlson pers. comm. 1991). Neither are there
any records of C. amoena from the “Top End” of
Australia, that is the northern coastline between
Cape York and the Kimberley coast.

The majority of records of Chelidonura
amoena now available come from Western Aus¬
tralia, but instead of indicating the species is
relatively more common on this coast than in
other parts of its range, the records may simply
reflect the greater number of opisthobranch ob¬
servers there than elsewhere in the range. Within
the continent of Australia, however, two biogeo-
graphical patterns are certainly evident. Cheli¬
donura amoena extends further south on the
west coast than on the east coast. On the west
coast, it extends to Rottnest Island (32°S),
whereas on the east coast it is only known from
Davies Reef (18°S) which is part of the northern
Great Barrier Reef. The greater influence of the
Leeuwin Current on the west coast than the East
Australian Current on the east coast probably
explains this anomaly. Secondly, C. amoena is
relatively much more abundant on the west coast
of Australia than on the east coast. Hundreds of
specimens have been observed along the north¬
west coast over the last decade (W. R. Rowlands,
G. McGarvie, C. W. Bryce pers. comm.), whereas
only a single individual has been discovered on
the east coast. At present we have no idea what
might be responsible for this imbalance. We
note, however, that the converse can also be true
for some species of Chelidonura. There is only
a single record of C.Julvipunctata from Western
Australia (Wells and Bryce 1993), yet the spe¬
cies is “common” along the east coast as far
south as Cape Byron (pers. obs.).

The opisthobranch Chelidonura amoena in Australia

ACKNOWLEDGEMENTS

Our foremost thanks are reserved for the
Underwater Scientific and Technical Research
Group of Florence for organising the Italian
Great Barrier Reef Expedition and thereby mak¬
ing Cattaneo-Vietti’s visit to Australia possible.
The discovery of Chelidonura amoena on that
expedition provided the impetus for this study.
Several colleagues contributed by supplying
information on C. amoena : Mr R. Moolenbeek
permitted us to examine the remaining type
material; Mr C. Carlson and Dr I. Hamatani
provided colour photographs of specimens from
the Palau Islands and Okinawa respectively;
Messrs C. W. Bryce, W. R. Rowlands and G.
McGarvie helped with records and photographs
from Western Australia; Mr McGarvie gave
permission to reproduce two of his colour slides.
Dr K. Baba confirmed the species’ absence from
Japan. Mr O. E. S. Kelly kindly translated the
original German description of C. amoena.

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Risbec, J. 1951. Notes sur les tectibranches de
Nouvelle-Caledonie. Journal du Society des
Oceanistes 7: 123-158.

Rudman, W. B. 1970. Chelidonura inornata Baba
and C. electro sp. nov. from the Solomon Islands
(Opisthobranchia, Aglajidae). Journal of the
Malacological Society of Australia 2: 7-12.

Rudman, W. B. 1973. On some species of Chelidonura
(Opisthobranchia : Aglajidae) from Zanzibar
and Fiji. Zoological Journal of the Linnean
Society 52: 201-215.

Rudman, W. B. 1974. A comparison of Chelidonura,
Navanax and Aglaja with other genera of the
Aglajidae (Opisthobranchia : Gastropoda). Zoo¬
logical Journal of the Linnean Society 54: 185-
212 .

Springsteen, F. J. and Leobrera, C. B. 1986. Sea-
shells of the Philippines. Carfel Seashell Mu¬
seum: Manila.

R.C. Willan and R. Cattaneo-Vietti

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Accepted 26 September, 1995

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:19-51

A REVIEW OF THE AUSTRALIAN ENDEMIC GOBIID FISH GENUS
CHLAMYDOGOBIUS, WITH DESCRIPTION OF FIVE NEW SPECIES.

HELEN K. LARSON

Museum and Art Gallery of the Northern Territory,
PO Box 4646, Darwin, NT 0801, Australia.

ABSTRACT

The Australian endemic gobiid genus Chlamydogobius is reviewed and five new
species described as new. One of the new species is estuarine, found in coastal northern
Australia; all the other species are restricted to freshwater springs and bores in inland
Queensland, South Australia and the Nonhem Territory. Chlamydogobius belongs to
the subfamily Gobionellinae and may be most closely related to Pseudogobius,
Hemigobius and Mugilogobius. An artificial key to species is given; species are
distinguished by morphology and vertebral counts. One of the new species is consid¬
ered to be endangered and two other new species are vulnerable.

Keywords: Chlamydogobius, Gobiidae, Gobionellinae, freshwater fish, estuarine fish,
new species.

INTRODUCTION

The desert goby, Chlamydogobius eremius
(Zietz, 1896), is probably one of the most well-
known gobiid fish species in Australia (Fig. 1). It
is widely kept by aquarium fanciers, and has
been referred to in a number of publications
dealing with inland fish fauna and ecology (for
example: Glover 1971; Glover 1973, Glover
1982; Glover 1989; Glover 1990; Glover and
Sim 1978;Horsthemke 1989). In recent years, it
has become apparent that some of the populations
of Chlamydogobius are very isolated and may be
endangered by increased pastoral activities and
reduction in the water table (Harris 1987; Wager
and Jackson 1993).

The taxonomic status of the genus and its
possible relationships were discussed by Miller
(1987), who considered it to be a monotypic
valid genus most closely related to Mugilogobius
mitt. Chlamydogobius is a gobionelline sensu
ezold (1993) and Larson (in prep.) and is here
considered to be most closely related to
Pseudogobius Popta, Hemigobius Bleeker and
Mugilogobius.

As Miller (1987) noted, Whitley (1930) erected
t e genus Chlamydogobius without any com¬

ment or diagnosis. Miller considered that, based
on the six species of Mugilogobius he had exam¬
ined, and information in Akihito et al. (1984),
Chlamydogobius should remain separate from
Mugilogobius due to i thaving a reduced number
of sensory papillae (especially row c), a longitu¬
dinal (versus transverse) row .v on the snout, a
single mental papilla on each side (versus a
transverse row or rows of papillae), reduced
number of scale ctenii, a much convoluted gut,
dark-pigmented peritoneum, one epural,
metapterygoid with spur not reaching across to
quadrate, and 28 vertebrae.

As will be shown in a revision of the
gobionellines related to and including
Mugilogobius, some of the above characters
given by Miller (1987) can be found among
some Mugilogobius species (Larson in prep.).
These characters include: reduced numbers of
mental papillae, black to brown peritoneum and
broad metapterygoid. Chlamydogobius has one
of the basic gobionelline characters, vertebral
formula 3-12210 ( sensu Birdsong et al. 1988),
and it shares with Mugilogobius the 9/7 seg¬
mented caudal ray pattern and basic sensory
papillae pattern (Fig. 1). Both genera have lost
all lateral line pores. The relationship of these

H.K. Larson

two genera and other similar gobionellines are
outlined by Larson (in prep.)

Harris (1987), Miller (1987), Glover (1989)
and Jackson (1993) have indicated that there
may be more than one “desert goby” species.
The author considers that there are six species in
this genus: five from freshwater desert habitats,
and one marine/estuarine. The marine species
had been recognised as undescribed by the au¬
thor for some time, but was only during the
course of this study that the relationship to C.
eremius became clear. Four of the freshwater
species and the marine species are described as
new below. A key to species is provided.

Chlamydogobius is easily distinguished from
any otherfish in the freshwaterenvironment, but
in the estuaries of northern Australia, one species
of Chlamydogobius can co-occur with species of
Hemigobius, Mugilogobius and Pseudogobius ,
which are similar in appearance and may be its
closest relatives (Larson: in prep.). Pseudogobius
has headpores in the interorbital region, a
rounded, somewhat inflated snout that usually
overhangs the upper lip and a long gut which
spirals in a corkscrew manner about itself along
its longitudinal axis. Hemigobius also has a few
interorbital headpores, the gut is long and coiled,
watch-spring-like, separate from the stomach,
but it has 17 segmented caudal rays, unlike the
other genera (which have 16 segmented caudal
rays). Mugilogobius is most similar to
Chlamydogobius, as it lacks headpores and has
similar basic external morphology; the status of
the two genera is currently under review. To
date, the two can be distinguished by the follow¬
ing characlei:s:Mugilogobius: s papillae on snout
in at least three rows of two or more papillae, the
first of which is usually the longest and runs just
above upper lip fold (few species with middle
row represented by only 1-2 papillae); pectoral
rays 13-20; intestine simple, with one “S-bend”
and no full loops; gill opening to pectoral base or
further, usually with fleshy knobs or ridge along
shoulder; two epurals, metapterygoid forming
distinct bridge to quadrate, 26-27 vertebrae, usu¬
ally 26 (10, rarely 11, precaudal and 16-17 cau¬
dal), males often with distinctly enlarged mouths.

Chlamydogobius : s papillae on snout usually
in two rows, rarely three (each row consists of only
1-2 papillae), first row just aboveupperlip absent;
pectoral rays 11-14; intestine long and coiled into
three loops separate from stomach; gill opening
restricted to pectoral base, shoulder girdle
smooth; one epural, metapterygoid not forming

bridge to quadrate, 27-29 vertebrae, usually 28
(10-11 precaudal and 16-18 caudal), males with
mouths not much larger than those of females.

Chlamydogobius is masculine; the exact deri¬
vation of the name is uncertain (Whitley gave no
etymology, as was his way). Whether Whitley
intended to refer to Chlamydes, nowadays a
junior synonym of Bathygobius (some species of
which, such as B. cotticeps, do resemble
Chlamydogobius in general appearance), or to
the more exact meaning of chlamydos (mantle or
cloak), is only speculation.

Methods. Measurements were taken using
electronic calipers and dissecting microscope.
Counts and methods generally follow Hubbs and
Lagler (1970), except as indicated below.
Pterygiophore formula follows Birdsong et al.
(1988). Transverse scale counts are taken by
counting the number of scale rows from the anal
fin origin diagonally upward and back toward
the second dorsal fin base. Head length is taken
to the upper attachment of the opercular mem¬
brane. Interorbital width is least fleshy width
(not least bony width). In the descriptions, an
asterisk indicates counts of the holotype (or
lectotype). Numbers in parentheses after counts
indicate the number of specimens with that count,
or the range of counts. Vertebral counts and
other osteological information was obtained by
radiography and clearing and double-staining.

Abbreviations: institutions - AMS, Australian
Museum, Sydney; NTM, Museum and Art Gal¬
lery of the Northern Territory, Darwin; QM,
Queensland Museum, Brisbane; ZMH,
Zoologische Museum, Hamburg; others - HD,
head depth at rear preopercular margin; HL,
head length; H W, head width at preopercle; my,
million years ago; SL, standard length; TRB,
transverse scale count backward.

Fig. 1 . Chlamydogobius eremius, papillae pattern. Scale
bar =■ 1 mm.

A review of the Australian endemic gobiid genus Chtamydogobius

SYSTEMATICS

Chtamydogobius Whitley, 1930

Chtamydogobius Whitley 1930: 122, central
Australia (type genus Gobius eremius Zietz,
1896, by original designation and monotypy).

Chalamydogobius - Lake 1971: 44 (lapsus).

Diagnosis. Genus distinguished by combina¬
tion of characters: single epural; 27-29 vertebrae
(10-11 precaudal and 16-18 caudal); usually one
more soft dorsal than anal ray present; long gut
with at least two full loops in the intestine, very
dark-pigmented peritoneum; gill-opening re¬
stricted to pectoral base; gill rakers without spines;
shoulder girdle smooth, without bony flange, or
fleshy bumps or flaps; jaws not much longer in
males compared with females; headpores ab¬
sent; row c on cheek below eye much reduced to
absent, .v rows on snout usually consist of one
(rarely two) papilla each, only two s rows present
in most species, usually only one/row mental
papilla on each side behind symphysis; male
breeding colour includes dark, white-margined
second dorsal, anal and caudal fins and yellow
band with blue spot below it on upper half of first
dorsal fin; genus restricted to the Australian
continent.

Osteology. Pterygiophore formula 3-12210;
27-29 vertebrae, 10-11 precaudal and 16-18 cau¬
dal vertebrae; neural spines of first thee verte¬
brae often stout, broadened or bifid at tip, usually
second and third spines bifid or broad-tipped;
usually two (occasionally one) anal fin
pterygiophores before haemal spine of first cau¬
dal vertebra; maxilla narrow, with one adductor
mandibulae attachment at half-way point; pala¬
tine broad anteriorly, becoming quite slender
ventrally, not quite reaching to quadrate and
sometimes falling well short of it; pterygoid
short, broad-based; metapterygoid broad, not
extended forward toward quadrate, with pointed,
broad-based flange or spur extending
anterodorsally; quadrate ratherforked, with lower
limb slender; symplectic without spur or process
extending toward preopercle; preopercle nar¬
row, without groove posteriorly; scapula
unossified; single epural, sometimesepural partly
split.

KEY TO SPECIES OF THE GENUS
CHLAMYDOGOBIUS

1 a. Second dorsal and anal fins usually with

equal number of rays, modally 1,6; pectoral
rays 11-12; usually 27 vertebrae; estua¬
rine, from northern coast of Australia....

..C. ranunculus n. sp.

b. Second dorsal fin with one more ray than
anal; second dorsal usually 1,7-8; anal usu¬
ally I, 6-7; pectoral rays usually 13-14 (if
11 -12, then second dorsal and anal not 1,6);
28-29 vertebrae; freshwater, restricted to
central Australia. 2

2 a. Opercles, pectoral base and breast usually

naked, predorsal region often naked; second
dorsal modally 1,8; anal modally 1,7; pec¬
toral usually 13; usually 28 vertebrae
(11+17); restricted to creeks and bores south

and west of Lake Eyre.C. eremius

b. Opercles, pectoral base and breast at least
partly scaled; predorsal region usually
scaled; second dorsal rays 1,6-8, anal rays
1,5-8; vertebrae 28-29.3

3 a. First dorsal fin reduced in size, spines III-

V (rarely VI); second dorsal 1,6-7; anal 1,5-

6; pectoral rays 11-12; 28 vertebrae; re¬
stricted to Elizabeth Springs, western
Queensland.C. micropterus n. sp.

b. First dorsal fin not reduced, spines usually

VI (sometimes V in gloveri n. sp.); second

dorsal 1,7-8; anal 1,6-7; pectoral usually
12-13; vertebrae 28-29.4

4 a. Cheek with small cycloid scales; usually

10 precaudal vertebrae; pectoral rays 13-
14, usually 13; restricted to springs around
Edgbaston Station, central Queensland

(Barcoo/Thomson drainage).

.C. squamigenus n. sp.

b. Cheek naked; usually 11 precaudal verte¬
brae; pectoral rays 11-14, usually 12 or 13;
from central Australia (NW of Lake Eyre)
.5

5 a. Second dorsal modally 1,7; anal modally

1,6; pectoral rays modally 12; usually 28
vertebrae (11+17); restricted to Dalhousie

Springs, northern South Australia.

.C. gloveri n. sp.

b. Second dorsal I, 7-8; anal modally 1,7;
pectoral raysmodally 13;usually 29verte-
brae (11+19); restricted to the upper Finke

River drainage, Northern Territory.

.C. japalpa n. sp.

H.K. Larson

Fig. 2. Lectotype of Gobius eremius, SAM F.525, 51 mm SL male.

Chlamydogobius eremius (Zietz, 1896)
(Figs 1-4, Plate 1, Tables 1-4)

Gobius eremius Zietz, 1896:180,pl. 16, fig. 5
(Central Australia). - McCulloch and Ogilby
1919: 257; - McCulloch 1929: 372.

Chlamydogobius eremius -Whitley 1930:122;
-Koumans 1931: 159-160;-Whitley 1964:123;

- Glover 1971: 1-147; - Glover 1973: 8-10;
Glover 1973: 4; Scott et al. 1974: 271-272; -
Glover and Sim 1978: 38; - Lake 1978: 73,153;
-Glover 1982: 242-244; - Thompson 1983: 17-
20; - Merrick and Schmida 1984: 309-310; -
Miller 1987: 687-705; - Glover 1989: 90-91;
Allen 1989: 203-204; - Horsthemke 1989: 288;

- Larson and Martin 1990: 62-63 (in part); -
Glover 1990: 189, 191, fig.Id.

Chalamydogobius eremius - Lake 1971: 44
(lapsus).

Material examined. 77 specimens (10-51).
SOUTH AUSTRALIA: lectotype of Gobius
eremius, SAM F.525,51 mm SL male. Coward
Springs, bore by railway, 4May 1894, coll. Horn
Expedition. Paralectotypes of Gobius eremius,
SAM F.7674,5( 13-43), same data as lectotype.
AMS 1.24673-001, 24(10-34.5), including two
cleared and stained, 6 km ESE of Coward Springs,
coll. D.F. Hoese and S. Reader, 23 August 1984;
SAM F.3509,10(30-44), Coward Springs Bore,
coll. J, Glover, 2 July 1968; SAM F.4204,6(27-
45.5), dam near Coward Springs, coll. J. Barry,
April 1976; SAM F.3999,13(32.5-41), Margaret
River, coll. T. Sim, 3 March 1975; AMS 1.26285-
001, 4(14.5-18.5), spring near Well Spring,
Freeling Springs, coll. W. Ponder, 2 June 1988;
AMS 1.24493-001, 4(28-41), including one
cleared and stained, Finniss Springs, coll. W.
Ponder, R. Hershler, 28 January 1984; AMS

1.27116-001, 10(13-29), Ockenden Spring and
Bore, coll. W. Ponder, 1 June 1983.

Other material examined (but not used in
description). Twenty-three specimens from the
following localities. AMS 1.30085-001, 2, Dav-
enportSprings,SA;AMSI.30087-001, ^.Eliza¬
beth Springs, SA; ZMH 31790, 2, aquarium
specimens, locality unknown; NTM S. 11426-
001,4, “The Bubbler”, mound springs, SA.

Diagnosis. A moderate to large Chlamydo¬
gobius with second dorsal rays 1,6-8; anal rays
1,5-7; pectoral rays 12-14; longitudinal scales
33-57 (usually 45-50); TRB 13-19; vertebrae
10+17 to 11+18 (mode 11+17); predorsal scales
0-10, scales, if present, scattered on nape midline,
arranged in rows on sides of nape only; most
scales cycloid, ctenoid scales on sides of body
extend in narrow wedge up to behind pectoral fin
or in twodistinct patches; opercles naked, pecto¬
ral base and breast usually naked, belly midline
sometimes naked; head and body light brown,
heavily mottled and marbled with darker brown
irregular markings (restricted to the western and
southern Lake Eyre drainage of South Aus¬
tralia).

Description. Based on 49 specimens, 14.5-51
mm SL. An asterisk indicates counts of the
lectotype (Fig. 2).

First dorsal IV (in 1), V (1), VI* (43); second
dorsal 1,6-1,8 (mean 1.8*); anal 1,5-8 (mean 1,7*,
1,6 in lectotype), pectoral rays 10-15 (mean 13*),
segmented caudal rays 16-17 (mean 16*); cau¬
dal ray pattern usually 9/7*; branched caudal
rays 14-16* (mean 15);unsegmented (procurrent)
caudal rays 7/6 (2), 7/7 (3), 7/8 (1), 7/9* (1), 8/
8 (1), 8/9 (1), 9/7 (1); longitudinal scale count
33-57 (mean 46, 44 in lectotype); TRB 13*-19
(mean 16); predorsal scale count 0* -10 (mean 6;

A review of the Australian endemic gobiid genus Chlamydogobius

0in41 specimens); circumpeduncular scales 16-
24 (mean 20, 19 in lectotype). Gill rakers on
outer face of first arch ranging from 2+5 to 3+10
(mode 2+6). Pterygiophore formula 3-12210*
(23). Vertebrae 10+17 (2), 11+16 (3), 11+17*
(26), 11+18 (3). Neural spines of first three
vertebrae variably split or expanded* at tip (16).
One* (28) or two epurals (3). One (3) or two*
(28) anal pterygiophores before haemal spine of
first caudal vertebra.

Body rounded anteriorly, compressed
posteriorly. Head short, somewhatrounded, wider
than deep, HL 2.8-3.5 (mean 3.2) times in SL;
cheeks may be inflated and fleshy in large males.
Depth at posterior preopercular margin 1.5-1.9
(mean 1.7) in HL. Width at posterior preopercular
margin 1.3-4.4 (mean 1.5) in HL. Mouth termi¬
nal to subterminal, slightly oblique, forming
angle of about 20° with body axis; upper jaw
slightly overhangs lower jaw; jaw length not
greatly different between males and females;
jaws generally reach to mid-eye in males and to
below front half of eye in females (to below mid¬
eye in lectotype). Lips usually fleshy, smooth,
usually without fleshy fimbriae visible on inner
edges (visible in lectotype, which is rather dehy¬
drated); lower lip free at sides, fused across front.
Upper jaw 2.1-3.7 (mean 3.2 in females, 2.8 in
males) in HL. Eyes lateral, high on head, top
usually forming part of dorsal profile, 2.6-5.6
(mean 4.4) in HL. Snout rounded to rather flat¬
tened, occasionally slightly inflated and partly
overhanging upperlip, 3.0-4.3 (mean 3.6) inHL.
Interorbital broad, flat to very slightly convex,
1.7-4.0 (mean 3.4) in HL. Top of head, from rear
of interorbital space up to snout tip, with scatter¬
ing of very fine villi (villi only visible in speci¬

mens with well-preserved mucous coat in this
area). Body depth at anal origin 4.6-5.8 (mean
5.3) in SL. Caudal peduncle compressed, rela¬
tively long, length 3.5-7.6 (mean 3.9) in SL.
Caudal peduncle depth 6.6-11.9 (mean7.9) inSL.

First dorsal fin low, rounded, second or third
spines longest or subequal; third spine usually
longest; spines do not reach second dorsal origin
when depressed. First dorsal spine always shorter
than next three. Second dorsal spine length 7.5-
10.0 (mean 9.1) in SL. Third dorsal spine length
6.6-10.6 (mean 8.8) in SL. Fourth dorsal spine
length 6.6-9.8 (mean 8.6) in SL. Second dorsal
and anal fins low, posteriormost rays longest,
when depressed, rays barely reach caudal fin in
large mature males; rays fall well short of caudal
fin in all other specimens. Pectoral fin broad,
rounded,central rays longest, 4.0-7.3 (mean 4.6)
in SL; rays usually all branched (lowermost ray
may be unbranched). Pelvic fins quite small,
rounded tooval, may reach up to half distance to
anus, 4.0-8.6 (mean 6.7) in SL. Caudal fin oval
to rounded or rectangular in form, 3.4-4.2 (mean
3.8) in SL.

No mental frenum, chin smooth. Anterior
nostril in short tube, placed on edge of preorbital,
tube oriented down and forward, preorbital some¬
times curved forward slightly to accommodate
nostril. Posterior nostril small, oval, placed half¬
way between front centre margin of eye and edge
of preorbital. Gill opening restricted to pectoral
base. Inneredge ofshouldergirdleeithersmooth
and fleshy, or hard-edged, with no bony flange
or fleshy knobs and flaps present. Gill rakers on
outer face of first arch very short rounded knobs,
without spines, longest one or two rakers near
angle of arch; outer rakers on second and third

Table 1, Measurements (mm) of Chlamydogobius eremius (Zietz, 1896).

Character

Lectotype

Minimum

Males

Maximum

Mean

Maximum

Females

Minimum

Mean

Head Length

17.0

6.0

17.0

10.8

6.7

12.4

9.6

Head Depth

9.0

3.8

9.0

6.5

3.7

7.8

5.6

Head Width

10.5

4.6

11.5

7.9

2.5

8.5

6.5

Body Depth

9.5

3.7

9.5

6.7

3.8

8.0

6.0

Body Width

1.8

5.9

3.9

2.5

5.2

3.7

Caud. Ped. Leng.

12.9

5.3

12.9

8.9

5.8

10.8

8.3

Caud. Ped. Depth

7.1

2.6

7.1

4.6

2.7

5.5

4.1

Snout

4.7

1.6

4.7

3.3

1.8

3.7

2.7

Eye

3.7

1.4

3.7

2.4

1.6

2.8

2.2

Jaw

7.1

1.9

7.4

4.2

2.0

4.1

3.0

Interorbit

5.3

1.8

5.3

.3.6

1.7

4.1

2.8

Pectoral

12.4

4.6

12.4

7.9

4.7

8.9

6.9

Pelvic

7.3

3.2

7.8

5.4

3.2

5.9

4.6

Caudal

5.8

13.2

9.1

5.7

9.9

8.2

Longest D1 spine

5.2

2.3

5.7

4.4

3.0

4.5

4.0

H.K. Larson

arches usually smaller than those on first arch,
outerrakers absent from fourth arch. Gill rakers
on inner face of first and all other arches nearly
twice the length of first arch inner rakers. Tongue
thick and fleshy, usually blunt to rounded. Teeth
in outermost row of upper jaw usually larger than
others, stout and curved or almost upright; be¬
hind outer row, three to five rows of slightly
smaller stout curved teeth; number of rows re¬
ducing to one or two at side of jaw (teeth in
lectotype, a large specimen, only slightly curved).
Teeth in lower jaw in five or six rows; teeth
arranged as in upper jaw apart from outermost
row being not much larger than those in rows
behind it. Teeth not much differing in size be¬
tween males and females.

Predorsal scales usually absent; when present,
midline of nape naked with scales extending
forward at sides to above opercle, or small scales
present immediately before first dorsal fin or
scattered unevenly over nape. Operculum gener¬
ally naked (seven specimens with few small
cycloid scales scattered on upper third to half;
usually only one to three scales present). Cheek
always naked. Pectoral base naked (one speci¬
men with few small cycloid scales present).
Prepelvic area usually naked, sometimes with
few small cycloid scales directly before pelvic
fin. Belly sometimes with naked mid-line, other¬
wise covered with cycloid scales and isolated
patch under pelvics of weakly ctenoid scales.
Ctenoid scales on side of body present at least as
scattered scales on caudal peduncle and patch
behind pectoral fin; peduncle scales may extend
forward in narrow wedge, often broken into
patch of ctenoid scales behind pectoral fin and
ctenoid scale wedge posteriorly.

Head pores absent as in all Chlamydogobius.

Sensory papillae pattern longitudinal, as in
Figure 1. Row p composed of many small papil¬
lae (characteristic of genus). Two s rows on
snout, rows consist of only one or two papillae.
F row composed only of two (occasionally three)
papillae.

Coloration of fresh material. Taken from
colour slides (Plate 1).

Females and non-breeding males. Head and
body greyish brown (mouse grey) with seven to
eight dull to dark brown saddles or patches
across back; saddles may be clearly defined
square blotches or highly marbled patches. Sad¬
dles do not extend past mid-side of body, here
replaced by series of brown marbled blotches,
square blotches or irregu lar spots, markings fade
ventrally very quickly to match background col¬
our on lower half of body. Top and sides of head
marbled and spotted with brown, one or two dark
brown bars from eye to upper lip may be present.
Lips almost bluish grey, with brownish grey
edges. Iris deep golden, with dark brown margin.
Silvery white peritoneum usually visible through
body wall (internal lining of body cavity black),
especially noticeable in gravid females. Base of
caudal with one or three dark brown to greyish
brown spots; spots may form Y-shape or vertical
bar at fin base, with lighter background colour
(yellowish brown) surrounding spots.

First dorsal fin greyish brown, with deep yel¬
lowish submarginal band, below this band, nar¬
row blue line widening posteriorly to form bright
blue spot. Second dorsal fin with light greyish
brown membrane, dark brown fin rays and darker
brown blotches along base. Caudal fin light
greyish brown with irregular, vertically oriented
rows of fine brown spots. Pectoral fin very pale
greyish brown to hyaline, fin base with brown
blotch on upper half. (Pelvics not visible on
available slides).

Breeding males. Dark brown saddles and
mottling on head and body diffuse to absent,
usually only about seven ill-defined square sad¬
dles across back. Body and top of head generally
plain greyish brown to yellowish brown, lower
half of head distinctly yellowish brown to golden
yellow (yellow increases ventrally). No distinct
bars or mottling discernible on head. Iris usually
pale gold, contrasting sharply with dull colour¬
ing ofhead.

Table 2. Frequency distribution of fin ray counts in Chlamydogobius species.

Species

Second dorsal
6 7

rays

Anal

rays

Pectoral rays

12 13 14 15

eremius n.sp.

3 1

gloveri n.sp.

japalpa n.sp.

micropterus n.sp.

ranunculus nsp.

squamigenus n.sp.

A review of the Australian endemic gobiid genus Chlamydogobius

First dorsal fin very dark grey, with yellow to
yellow ochre submarginal band; below this band,
bright iridescent light to vivid blue spot on pos¬
terior half of fin. Second dorsal and anal fins
similar in colour to each other: both fins very
dark grey to blackish with relatively broad bright
white margin; basal half of fins speckled with
vivid to dark blue, blue often more distinct on
membrane between fin rays. Caudal fin similar
to second dorsal and anal fins, but with only
narrow dull whitish fin margin (margin not bright
white), and very little blue present. Pectoral fin
with membranes translucent to light dusky, fin
rays greyish. Pelvic fin translucent tolight dusky.

Coloration of preserved material. Colour
not greatly differing from live fish, but blue and
yellow pigment absent. Specimens vary in
amount and kind of marbling and spotting. Mar¬
bling more distinct in preserved material; lower
mid-side blotches often visible as series of elon¬
gate brown blotches ending in Y-shaped caudal
blotch (this blotch often much more distinct in
preserved than live specimens). On head, usu¬
ally one brown bar from front of eye to mid upper
lip and one brown bar from lower edge eye to just
behind rictus, with conspicuous pale interspace
between bars. Pectoral base pale with brown
blotch on upperhalf. Entire lining of body cavity
dense black.

Comparisons. Chlamydogobius eremius is
most similar to C.japalpa n. sp. Although fin ray
counts are similar, the extent of the scalation
differs between the two (predorsal n aked or with
sides of nape scaled and scattered scales on
midline in C. eremius ; predorsal, including
midline, usually scaled over opercle or to above
preopercle in C. japalpa n. sp.; pectoral base,
part of breast and opercle usually with some
scales in C. japalpa n. sp., with these areas
usually naked in C. eremius ) and C. eremius
modally has 17 caudal vertebrae (rarely 18,
which is the mode for C. japalpa n. sp.) and
never 19 as has been recorded in C. japalpa n. sp.
There are modal differences in second dorsal fin
ray and transverse scale counts and mean and

Table 3. Frequency distribution of transverse backward
scale counts in Chlamydogobius species.

Species 11

eremius n.sp.

gloveri n.sp. 1

japalpa n.sp.

1 1

micropterus n.sp.

ranunculus n.sp.

squamigenus n.sp. -

modal differences in lateral line scales (Tables 2-
4).

Mark Adams (pers. comm.) found that these
two species are very close electrophoretically
(differing by one locus) and considers that they
may not warrant separation as species. The Finke
River apparently has not discharged into Lake
Eyre during recorded history (Kotwicki 1989)
and may not have done so for 10,000 years or
more (possibly not since the last glacial maxi¬
mum of about 18,000 years).

Distribution. Specimens are known from the
southern and western Lake Eyre drainage of South
Australia. Localities (Fig. 13) from which the
species has been recorded are; Forrest’s
Waterhole, Ockenden Spring, Algebuckina
Waterhole, Wood Duck Bore, Peake Creek Sid¬
ing, Old Peake Homestead Bore, Freeling
Springs, Blythe Bore, Birribirriana Spring,
N ilpi nna Spring, Fi nniss Spri ngs, North Fou nt ai n
Spring, Johnson’s Number 3 Bore, Honeymoon
Bore, Nunn’s Bore, Wobna Spring, Strangways
Springs Railway Bore, Warriner’s Creek,
Beresford Reservoir, Railway Bore at Coward
Springs, Coward Springs proper, Chamber’s
Creek, MargaretCreek, Wobna Spring, Elizabeth
Springs, Anna Spring East Bore, un-named spring
SE of Wobna Spring, Davenport Springs, Dead
Woman Springs, Gregory Creek, Alberrie Creek,
Clayton’s Bore, Montecolina Bore and Woolatchi
Bore. Glover (1971) gives latitudes and
longitudes for these localities as well as for many
other springs not inhabited by fish.

The last two mentioned localities, near Lake
Blanche and Lake Callabonna respectively, ap-

Table 4. Frequency distribution of lateral line counts in Chlamydogobius species.

Species 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57

eremius n.sp. - 1 ... 1 1 13241446232441 1 - 1 - 1

gloveri n.sp. ---5-16511 1 ----- -.- - -

japalpa n.sp. ....1-12453-6414----- .

microplerus n.sp. - - - 1 3 1 3 2 4 4 2 3 - - - 1 .

ranunculus n.sp. 212-12374475531222--1 .

squamigenus n.sp. -1332263321132 .

H.K. Larson

parentlyhad fish in them in 1969 (Glover 1971)
but Glover failed to find any during his 1968-69
survey. Additionally, Glover (1971) introduced
50 male and 50 female adult C. eremius into
Blanche Cup Mound Spring, in September 1970,
in an experiment to determine the size and age
structure of a wild-living population; the intro¬
duced population flourished.

Zietz’s (1896) original description states that
his specimens were found in a small pool of
water aroundan artesian well at Coward Springs.
There has been some confusion over this, as
there is now more than one bore sunk in the
vicinity of these springs. In the narrative of the
Horn Expedition, Spencer (1896) states that "By
the railway side at Coward Springs one of these
[artesian bores] has been sunk...", confirming
Glover’s (1973; unpublished notes) indications
that the Coward Springs Railway Bore (29° 24’S
136° 49’E) is the type locality (and not Coward
Springs proper).

Ecology. Glover’s publications (Glover 1971,
1973, 1982, 1990; Glover and Sim 1978) give
considerable detail on this adaptable species’
physiology,environmental tolerances, dispersal
and feeding behaviour, which will not be re¬
peated here.

Chlamydogobius eremius has been shown, by
experimentation in the field and the laboratory,
to be tolerant of a considerable range of tempera¬
ture, pH and salinity (Glover 1971; Glover and
Sim 1978). It occurs in waterof pH 6.8-11.0, can
withstand a temperature range of 5-41°C and
salinity range of 0-60 %o for short periods, and
has been collected in water with very low levels
of dissolved oxygen (0.8 ppm), frequently in¬
habiting water with concentrations below 5 mg
O/l (Glover 1971; 1982).

Captive maintenance, behaviour and spawn¬
ing have been reported by Thompson (1983),
Horsthemke (1989), Wilson (1992) and in many
other “aquarium hobby” magazines. It is not

Fig. 3. Distribution of the genus Chlamydogobius. * = C. eremius. ■ = C. gloverin. sp., V = C.japalpa n. sp., A = C. micro
pterus n. sp., • = C. ranunculus n.sp., D =C. squamigenus n. sp.

A review of the Australian endemic gobiid genus Chlamydogobius

Fig. 4. Known localities of the freshwater species of Chlamydogobius (n.b. the two easternmost populations of C. eremius

H.K. Larson

certain that the fish reported on were all C.
eremius, as description and provenance of the
captive populations were not always given or
wereunclear (Thompson clearly stateshis speci¬
mens were collected from Coward Springs, the
type locality of C. eremius , while Wilson’s speci¬
men's may have been C.japalpa n. sp.). Glover
(1971) was not able to provide conditions suit¬
able for his specimens to spawn.

Horsthemke (1989) gives detailed descrip¬
tions of embryonic and larval development, and
noted that there was no planktonic larval stage;
the hatchlings (5.3 mm long) quickly (precise
time not given) taking up a benthic existence. He
reported that up to 300 eggs may be laid by large
females, while Glover (1971) found 150-250
eggs in the ovaries of wild-caught specimens.
Miller (1987) correctly estimated that newly
hatched Chlamydogobius larvae were likely to
be “relatively large, probably about 4-5 mm”
and compared egg sizes and numbers for C.
eremius. Mugilogobius abei and Pseudogobius
olorum. Horsthemke (1989) compared spawn¬
ing and the direct larval development of C.
eremius with that of M. abei and M. chulae (the
latter two Mugilogobius have a four to five week
planktonic larval life).

Michaelis (1985) listed this species as not
threatened, along with a number of other fresh
and brackish water fishes with poorly known
distributions and abundances.

Remarks. The largest male, possibly the speci¬
men illustrated by Zietz (1896), is hereby desig¬
nated lectotype of the species Gobius eremius
(Fig. 2). Spencer (1896) refers to C. eremius
occurring around the opening of the artesian
bores at Coward and Strangways Springs, and
states that the Coward Springs bore "... issues at
a temperature of 95° F\ He considered it to be of
“little doubt” that the two localities were stocked
by fish hatched from eggs attached to feet or
feathers of birds, although Zietz (1896) only
stated that it was “possible”.

Chlamydogobius gloveri n. sp.

(Figs 3-6, Plates 1-2, Tables 2-5)

Chlamydogobius eremius - Glover 1971: 77,
99, Table 1, Appendix A; - Ivantsoff and Glover
1974: 95.

Chlamydogobius sp. nov. (Dalhousie goby) -
Glover 1989: 90, fig.31.lg; - Glover 1990: 191.

Chlamydogobius n. sp. - Harris 1987: 9;
Jackson 1993: 24.

Chlamydogobius sp. - Kodric-Brown and
Brown 1993: 1850; - Morton etal. 1995:30,95.

Material Examined. 23 specimens (13-36).
DALHOUSIE SPRINGS, SOUTH AUS¬
TRALIA: HOLOTYPE - SAM F.3463, 30 mm
female, coll. J. Glover, August 1968.
PARATYPES - SAM F.5425, 12(13-27.5),
Spring “G”ab, coll. J. Glover and T. Sim, 4 June
1985; SAM F.7675, 3(33-36), same data as |
holotype; SAM F.5417, 4(18-24), Spring B4,
coll. J. Glover, T. Sim, 3 June 1985; AMS
1.27118-001, 1(32), pool on top of mound, low
mound springs, coll. W. Ponder, 29 May 1983;
AMS 1.25881-001,4(7-16), Cold Spring Cc 1B,
outflow of medium active spring, coll. W. Pon¬
der, D. Winn, 6 June 1985; AMS 1.25879-001,
1(20), warm pool in swamp, coll. W. Ponder, D.
Winn, 13 June 1985; AMS 1.25880-001, 1(18),
warm pool 20 m upstream from main pool, coll.
W. Ponder, D. Winn, 14 June 1985.

Other material examined (but not used in
description). Nine specimens from Dalhousie
Springs: AMS 1.25883-001, 1; AMS 1.25882-
001,1; AMS 1.25876-001,1; AMS 1.25867-001,

1; AMS 1.25884-001, 1; AMS 1.25877-001, 1.

Diagnosis. A small Chlamydogobius with sec¬
ond dorsal rays 1,6-8; anal rays 1,6-7; pectoral
rays 11-13; longitudinal scales 35-43; TRB 11-
16; predorsal scales 13-18, small, reaching to
above preopercular margin or up to behind eyes;
most scales ctenoid, ctenoid scales present in
patch behind pectoral fin, under pelvic fins, and
variably on caudal peduncle; dorsal fin low;
brown with darker mottling which may form one
or more lateral stripes; known only from
Dalhousie Springs, northern South Australia.

Description. Based on 20 specimens, 15.5-36
mm SL. An asterisk indicates counts of holotype
(Fig. 5).

First dorsal V (4), VI* (15); second dorsal 1,6-
8 (mean 1,7*); anal 1,6-7 (mean 1,6*), pectoral
rays 11-13 (mean 12*), segmented caudal rays
15-17 (mean 16*); caudal ray pattern usually 8/

7* or 9/7; branched caudal rays 13-16 (mean
15*); unsegmented (procurrent) caudal rays 7/6
(2); longitudinal scale count 35-42 (mean 38);
TRB 11-16 (mean 14*); predorsal scale count
13-18(mean 17*), circumpeduncular scales 16-
19* (mean 18). Gill rakers on outer face of first
arch ranging from 2+6 to 3+7 (mode 2+7*);
pterygiophore formula 3-12210 (7). Vertebrae
10+18(2), 11+16(1), 11+17(10).Neural spines
of first few vertebrae slender, narrow (3). One
epural (7), possible twoepurals in one specimen

A review of the Australian endemic gobiid genus Chlamydogobius

Fig. 5. Holotype of Chlamydogobius gloveri n. sp., SAM F.3463, female.

(difficult to discern in X-ray). One (2) or two (6)
anal pterygiophores before haemal spine of first
caudal vertebra.

Body rounded to compressed anteriorly, com¬
pressed posteriorly. Head wider than deep, but
not greatly so, HL 3.1-3.6 (mean 3.42) in SL;
head rounded to somewhat rectangular in cross-
section.. Depth at posterior preopercular margin
1.5-1.8 (mean 1.7) in HL. Width at posterior
preopercular margin 1.2-1.6 (mean 1.5) in HL.
Mouth terminal, rarely slightly subterminal,
slightly oblique, forming angle of about 25° with
body axis; jaws small, generally reach to below
front half of eye. Lips usually smooth, fleshy
fimbriae sometimes present on inner edges of
upper lip; lower lip free at sides, fused across
front. Upper jaw length not much differing be¬
tween males and females, 3.0-3.5 (mean 3.1 in
females, 3.2 in males) in HL. Eyes lateral, high
on head, top usually forming part of dorsal

profile, 3.5-4.7 (mean 3.9) in HL. Snoutrounded,
sometimes lightly inflated but not overhanging
upper lip, 2.8-4.4 (mean 3.7) in HL. Interorbital
moderately broad,flat, 3.0-3.9 (mean 3.5) in HL.
Unsealed portion of top of head rarely with fine
villi under mucous coat. Body depth at anal
origin 4.5-5.8 (mean 5.2) in SL.Caudal peduncle
compressed, length 3.3-3.9 (mean 3.6) in SL.
Caudal peduncle depth 6.8-8.0 (mean 7.4) in SL.

First dorsal fin quite low, rounded, third or
fourth spines longest or subequal; spine length
not much different between males and females;
spines fall short of second dorsal fin origin when
depressed. First and second dorsal spines always
shorter than next two. Third dorsal spine length
8.3-11.3 (mean 9.7) in SL. Fourth dorsal spine
length 8.3-10.7 (mean 9.5) in SL. Second dorsal
and anal fins short-based, very low, posteriormost
rays usually longer than others, rays fall well
short of caudal fin base when depressed. Pectoral

Table 5. Measurements (mm) of Chlamydogobius gloveri n. sp.

Character

Holotype

Minimum

Males

Maximum

Mean

Maximum

Females

Minimum

Mean

Head Length

8.3

5.0

8.1

6.3

6.1

10.5

8.2

Head Depth

5.6

2.9

5.1

3.8

3.7

6.6

5.2

Head Width

6.5

3.4

5.7

4.4

4.1

8.3

6.1

Body Depth

6.6

2.8

5.3

3.9

4.1

7.7

6.0

Body Width

4.0

1.5

3.5

2.5

2.3

4.5

3.8

Caud. Ped. Leng.

8.7

4.5

7.6

5.8

10.2

8.0

Caud. Ped. Depth

4.3

2.0

3.8

2.8

2.7

5.1

3.9

Snout

2.7

1.2

1.8

1.6

1.7

3.7

2.5

Eye

2.0

1.3

2.0

1.7

1.5

2.7

2.0

Jaw

2.8

1.5

2.6

2.0

1.9

3.4

2.7

Interorbit

2.5

1.3

2.6

1.9

1.7

3.3

2.5

Pectoral

6.1

3.7

6.1

4.6

4.0

7.4

5.7

Pelvic

4.4

2.1

4.2

3.2

3.3

5.5

4.3

Caudal

7.1

4.7

6.8

5.7

5.6

8.4

7.1

Longest D1 spine

2.8

2.7

3.1

2.9

2.8

3.3

3.1

H.K. Larson

fin rounded, central rays longest, 4.3-5.5 (mean
4.7) in SL; rays usually all branched. Pelvic fins
short, rounded tooval, may reach half distance to
anus, 6.0-7.6 (mean 6.6) in SL. Caudal fin
rounded, 3.4-4.3 (mean 3.8) in SL.

No mental frenum, chin smooth. Anterior
nostril in very short tube on preorbital just be¬
hind upper lip, tube oriented down and forward,
preorbital curved forward slightly to accommo¬
date nos tri 1. Posterior nostri 1 small, round, placed
about halfway between front margin of eye and
edge of preorbital. Gill opening restricted to
pectoral base. Inner edge of shoulder girdle
smooth with no bony ridge or fleshy knobs or
flaps. Gill rakers on outer face of first arch very
short, unspined rudimentary knobs, longest raker
near angle of arch; rakers on inner face of first
arch also short but longer than outer rakers; inner
rakerson other arches nearly twice length offirst
arch innerrakers. Tongue broad, usually blunt or
rounded. Teeth slightly larger in male compared
to female. Outermost teeth in upper jaw arranged
in even row, teeth curved, longer than inner
teeth; behind this row are only one or two rows
of quite small sharp teeth; rows narrow to one at
side of jaw; tips of teeth sometimes lightly tinted
translucent brown. Lower jaw with about three
rows of small, curved, pointed teeth across front,
outermost row may be larger, stouter or less
curved; usually only one row at side of jaw; teeth
sometimes tinted translucent brown.

Predorsal scales small, evenly sized, usually
reaching forward to above preopercular margin
or further up to behind eyes. Operculum with
patch of small cycloid scales on upper three-
quarters to half. Cheek always naked. Pectoral
base covered with cycloid scales. Prepelvic area
covered with small cycloid scales. Belly with
isolated patch of ctenoid scales under pelvics,
rest cycloid. Most body scales cycloid, ctenoid
scales on side of body present as patch behind
pectoral fin, variably present along mid-line of
body from posterior caudal peduncle forward;
posterior ctenoid scales usually not reaching
ctenoid patch behind pectoral fin.

Head pores absent as in all Chlamydogobius.

Sensory papillae pattern longitudinal, as in
Fig. 6. Cheek row c reduced, broken; rear portion
close to front part of row b. Two or three 5 rows
of snout; posteriormost row often of two papil¬
lae, central row (if present) consists of one pa¬
pilla; s rows placed well back from upper lip.
Mental/row consists of two papillae.

Coloration of fresh material. Taken from
colourslides. Male and female similar,except fin

colour more pronounced and body pattern darker
and somewhat obscured in male (Plate 1-2).

Head and upper sides of body bluish grey
greenish grey; lower sides paler, peritoneurq
whitish, showing through body wall. Top Qf
head and at least upper half of body covered with
fine greyish brown vermiculations and spotting;
most scales with greyish brown margin or spot
present, brown markings may form line along
mid-side of body, especially noticeable
posteriorly. Snout, suborbital, upper half of
preopercle and opercle with indistinct greenish
brown mottling, lowerhalfofopercle plain whit¬
ish. Lips greenish grey, lower lip paler than
upper lip in male.

First dorsal fin (in female) mostly translucent
with white blotches proximally, and scattered
blue spots forming central band, posteriormost
blue spot largest. In male, first dorsal fin light
golden brown proximally, with broad light blue
band across middle of fin, band narrow anteriorly,
widening posteriorly to occupy most of rear of
fin; uppermost third of fin dull yellow. Second
dorsal fin (in male) dull olive brown, with sub¬
marginal broad brownish grey band; lower half
of fin with scattered light blue and golden spots;
golden spot more abundant on lower half of fin.
Seconddorsal fin (in female) translucent brown¬
ish with pale gold to whitish spots and streaks
alternating with light brown spots; anteriormost
quarter of fin plain translucent.

Caudal fin translucent brownish with many
vertical rows of fine light brown to dull whitish
spots and small blotches; markings more diffuse
on ventral part of fin. Anal fin translucent brown¬
ish with dull yellowish white or blue marginal
band and scattered yellowish white or blue
patches or streaks on proximal third of fin. Pec¬
toral fin translucent, rays dusky, especially proxi¬
mally. pelvic fins whitish to translucent brown.

Fig. 6. Chlamydogobius gloveri n. sp., papillae pattern.
Scale bar = 1 mm.

A review of the Australian endemic gobiid genus Chlamydogobius

Coloration of preserved material. Head and
body light brown or yellowish white, with black¬
ish, brown to brownish grey irregular blotches
and spots (depending upon state of preserva¬
tion). Most conspicuous markings are: one series
of about six irregular brown blotches across
dorsal mid-line and second series of about seven
elongate brown blotches along mid-line of body,
anteriormost of which is just above or behind
pectoral base, posteriormost is at centre of cau¬
dal base (posteriormost spot often darker than
others). Blotches across dorsal mid-line often
broken into two-part linear blotches: one part on
dorsal mid-line and one part on upper middle of
body; latter series of markings tends to form
discontinuous streak. Pectoral base whitish on
lower half; brownish with dark brown spot on
upper half. Top of head above rear edge of
preopercle with brownish band or saddle; top of
snout with indistinct brownish vermiculations
and spots. Side of head dark on dorsal half, pale
ventrally; brown bar from front of eye to preorbital
edge, indistinct brown bar from ventral edge of
eye to just below rictus; oblique brown bar or
blotch across rear of preopercle.

First dorsal fin with greyish brown margin,
whitish to dusky grey band below this; proximal
half of fin dusky or mottled with brown, with
blackish to dense black spot at rear of fin (spot
variable in size). Second dorsal fin with narrow
whitish margin; remainder of fin mottled with
dusky brown. Anal fin with narrow whitish mar¬
gin; most of fin whitish with indistinct dusky
mottling proximally. Caudal fin translucent to
whitish, with up to nine vertically oriented rows
of brown spots, spots often joined together to
form irregular wavy lines; very narrow whitish
margin sometimes present; indistinct pairofelon¬
gate brown blotches on either side of centre of
caudal base, close to posteriormost mid-lateral
spot. Pectoral fin with dusky to brown rays,
membranes translucent whitish to dusky. Pelvic
fins whitish, frenum and ray bases often dusky.

No large males in breeding colour pattern
among material examined. Figure 13.1 g in Glover
(1989), although small, clearly shows the two
dark spots at the base of the caudal fin, and the
rows of dark spots on the fin.

Comparisons. Morphologically, C. gloveri is
similar to C. japalpa n. sp. (both species have
scaled napes) but usually C. gloveri has 17 cau¬
dal vertebrae (16-18, modally 17; versus 17-19,
modally 18), second dorsal rays 1,7 (versus 1,8),
anal rays 1,6 (versus 1,7) and only 12 pectoral
rays (versus 13).

Distribution. Specimens are only known from
spring-fed pools at Dalhousie Springs (26° 28’S
135° 29’E), in northern South Australia, where
they have been reported as being “very common
though variable in abundance” (Glover 1990).
Glover (1989) records C. gloveri as being present
in 30 of the 35 springs inhabited by fish (there are
89 springs in total).

Ecology. Glover (1989) gives a detailed ac¬
count of the physical conditions at Dalhousie
Springs and the composition of the fish species
assemblages present. The springs are warm (21.6-
43°C), and the gobies apparently have a thermal
tolerance of up to 43.9°C. Glover (1989) ob¬
served that Chlamydogobius gloveri' s diet in¬
cluded filamentous green algae and their own
species. Ivantsoff and Glover (1974) and Crowley
and Ivantsoff (1990a), in their respective de¬
scriptions of Craterocephalus dalhousiensis and
Craterocephalus gloveri, give additional envi¬
ronmental data (including colour photographs of
the habitat in Crowley and Ivantsoff).

Kodric-Brown and Brown (1993) analysed
fish community structure in 43 springs at
Dalbhousie, and found that the communities
exhibited "... an amazingly regular, determinis¬
tic structure: the number of species is highly
correlated with spring size ...". They found that
C. gloveri has apparently become extinct in one
very small spring (E3) since 1989 (Kodric-Brown
and Brown 1993: 1853).

Remarks. Glover (1989: 110-111) reported
that isozyme electrophoretic analyses of the heads
of the Dalhousie goby (C. gloveri), Elizabeth
Springs goby (C. micropterus n. sp.) and C.
eremius indicated that the three species show,
relative to C. gloveri and themselves, percent¬
ages of fixed differences of 21% and 16% re¬
spectively. It is not surprising that the lower
percentage separates C. gloveri and C. eremius,
as they are closer together (geographically).
Glover suggested that the Dalhousie goby and
the Elizabeth Springs goby are distinct species,
but did not discuss the Nilpinna Spring popula¬
tion, for which he gave a fixed difference per¬
centage of 18%. Nilpinna Spring is part of the
Nilpinna Creek system just north-west of Lake
Eyre, and is inhabited by C. eremius (Glover
1971; this study). Crowley and Ivantsoff (1990a:
119) state that the Dalhousie Springs have been
isolated for 10,000 years and that floodwaters
from the Eyre/Finke system have not reached the
springs since then. This is based on Kotwicki
(1989), mentioned earlier, who discussed the
isolation of the Dalhousie Springs region from

H.K. Larson

the rest of the Lake Eyre drainage basin, and
suggested that the Finke does not flow near
Dalhousie during flood events (that it may have
done so during the period of about 45,000 and
25,000 years before present is more likely).

Jackson (1993) lists this species as having
“restricted” conservation status. The true status
of this species should be investigated, along with
that of the other freshwater species of this genus,
as indicated by Glover (1990). The conservation
status of this species was first noted in Harris
(1987), where it was listed as “Restricted” (ie.
having a restricted distribution but not yet at risk).

Etymology. Named for the late John Glover,
in recognition of the considerable workhe carried
out on desert gobies and other Australi an arid zone
fishes. John was convinced that the Dalhousie
goby was separate from C. eremius ; he just never
quite got around to describing it. It is somehow
appropriate that there should be more than one fish
speciescalled gloveri living at Dalhousie Springs
{Craterocephalus gloveri was described by Lucy
Crowley and Walter Ivantsoff in 1990, and the
undescribed Dalhousie Neosilurus is rumoured to
become named after John also).

Chlamydogobius japalpa n. sp.

(Figs 3-4,7-8, Tables 2-4, 6)

Chlamydogobius eremius - Larson and Martin
1990: 62-63 (in part).

Material examined. 84 specimens (12-44).
FINKE RIVER, NORTHERN TERRITORY:
HOLOTYPE - NTM S.l 1436-007, 44 mm SL
male, Ormiston Creek, at junction of Pioneer
Creek, 23° 40’S 132° 42’E, coll. H. Larson and
P. Homer, 15 September 1984. PARATYPES -
NTM S.l 1436-009, 41(15-44), same data as

holotype; NTM S.l 1439-006, 10(15-38.5), in¬
cluding one cleared and stained, off Palm Valley
Road just N of Park border, coll. H. Larson, 17
September 1984; AMS 1.35467-001, 8(15-28),
same data as preceding; NTM S.l 1437-005,
25(12-45), just above Glen Helen N of main road
crossing, coll. H. Larson and P. Homer, 16
September 1984; NTM S.l 1639-001, 1(40),
Hermannsburg Rockhole, coll. R. Moses, 21
June 1983; SAM F.7677,3(25-34), same data as
preceding; NTM S. 11628-001, 1(29.5), Boggy
Hole, coll. R. Moses, 23 June 1983; NTM
S.l 1650-002,2(25.5-27), Running Waters, coll.
R. Moses, 14 July 1983; NTM S.l 1632-001,
1(37), Palm Valley, 25 June 1983.

Other material examined (but not used in
description). Eight specimens from the follow¬
ing localities. NTM S. 11651 -001, Running Wa¬
ters, Finke River; NTM S.l 1653-003, same lo¬
cality; NTM S.l2503-001, Finke River pools
near highway; NTM S.1667, Hermannsburg.

Diagnosis. A moderate to large Chlamydo¬
gobius with second dorsal rays 1,7-8; anal rays
1,5-8; pectoral rays 12-14; longitudinal scales
36-47; TRB 14-19; 28-29 vertebrae; predorsal
scales 7-18, scales nearly always present on mid¬
line of nape, reaching forward over opercle, or to
above preopercular margin; pectoral base and
opercle often with few scattered scales; ctenoid
scales present on caudal peduncle, sometimes
forming narrow wedge forward to behind pecto¬
ral fin; dorsal fin low; head and body brown with
darker spots and mottling which may form lat¬
eral stripe; known only from Finke River system
in central Northern Territory.

Description. Based on 32 specimens, 18.5-44
mm SL. An asterisk indicates counts of the
holotype (Fig. 7).

Fig. 7. Holotype of Chlamydogobius japalpa n. sp., NTM S.l 1436-007, male.

A review of the Australian endemic gobiid genus Chlamydogobius

First dorsal V (in 3), VI* (28); second dorsal
I,7*-I,8 (mean 1,8); anal 1,5-8 (mean 1,7; 1,6 in
holotype), pectoral rays 12-14 (mean 13*), seg¬
mented caudal rays 16; caudal ray pattern usu¬
ally 8/7* or 9/7; branched caudal rays 15-16
(mean 15*); unsegmented (procurrent) caudal
rays 7/7 (2); longitudinal scale count 36-47 (mean
43,45 in holotype); TRB 14-19 (mean 17,15 in
holotype); predorsal scale count 7-18 (mean 14,

11 in holotype); circumpeduncular scales 15-22
(mean 19, 18 in holotype). Gill rakers on outer
face of first arch ranging from 2+5* to 3+6
(mode 2+5); pterygiophore formula 3-12210
(15). Vertebrae 10+18 (3), 10+19(4), 11+17(2),
11+18(12). Neural spinesof first three vertebrae
expanded at tip (10) or stout and pointed (1). One
epural (17). One (1), two (16) or three (1) anal
pterygiophores before haemal spine of first cau¬
dal vertebra.

Body rounded to slightly compressed
anteriorly; compressed posteriorly. Head wider
than deep, HL 3.2-3.7 (mean 3.4) in SL; cheeks
may be quite fat in mature males. Depth at
posterior preopercular margin 1.6-1.9 (mean 1.8)
in HL. Width at posterior preopercular margin
1.2-1.5 (mean 1.4) in HL. Mouth terminal to
subterminal, slightly oblique, forming angle of
about 20° with body axis; upper jaw slightly
overhanging lower jaw; jaws generally reach to
below mid-eye in males and to below front half
of eye in females (to below mid-eye in holotype).
Lips usually smooth, fleshy fimbriae sometimes
visible on innermost edges of upper lip; lower lip
free at sides, fused across front. Upper jaw 2.2-
4.8 (mean 3.2 in females, 3.0 in males) in HL.
Eyes lateral, high on head, top usually forming
part of dorsal profile, 3.0-5.1 (mean 4.0) in HL.

Snout rounded to somewhat inflated and partly
overhanging upper lip, 2.1-4.0 (mean 3.5) in HL.
Interorbital broad, flat, 1.3-4.3 (mean 3.3) in HL.
Top of head, from above preopercular margin up
to snout tip, often with very fine pigmented villi
(in many recently collected specimens, villi and
surrounding mucous coat well-preserved). Body
depth at anal origin 4.5-5.9 (mean 5.1) in SL.
Caudal peduncle long, compressed, length 3.1-

7.5 (mean 3.7) in SL. Caudal peduncle depth 3.7-

8.5 (mean 7.4) in SL.

First dorsal fin low, rounded, third or fourth
spines longest orsubequal; in large males, spines
barely reach second dorsal origin when depressed;
spines fall well short of second dorsal fin in all
other specimens. First dorsal spine always shorter
than next three. Second dorsal spine length 8.5-

9.6 (mean 9.1) in SL. Third dorsal spine length
7.5-10.7 (mean 9.2) in SL. Fourth dorsal spine
length 7.3-10.3 (mean 9.1) in SL. Second dorsal
and anal fins low, short-based, posteriormost
rays longest, but rays fall well short of caudal fin
base when depressed. Pectoral fin rounded, cen¬
tral rays longest, 4.3-9.6 (mean 4.9) in SL; rays
usually all branched. Pelvic fins quite short, oval
to somewhat pointed (innermost rays longest),
may reach only half distance (or less) to anus,
5.0-7.1 (mean 6.6) in SL. Caudal fin oval to
rectangular in form, rounded posteriorly, 3.1 -6.9
(mean 3.8) in SL.

No mental frenum, chin smooth. Anterior
nostril in very short tube, placed on edge of
preorbital, tube oriented down and forward,
preorbital curved forward slightly to accommo¬
date nostril. Posterior nostril oval, placed half¬
way between front edge of eye and edge of
preorbital. Gill opening restricted to pectoral

Table 6. Measurements (mm) of Chlamydogobius japalpa n. sp.

Character

Holotype

Minimum

Males

Maximum

Mean

Maximum

Females

Minimum

Mean

Head Length

13.5

5.6

13.5

9.4

5.8

12.0

8.9

Head Depth

8.3

2.9

8.3

5.4

3.1

7.5

5.1

Head Width

11.2

3.7

11.2

6.9

3.8

9.1

6.4

Body Depth

9.4

3.6

9.3

6.2

3.4

9.3

6.3

Body Width

5.9

2.2

5.9

4.0

2.1

5.5

4.1

Caud. Ped. Leng.

11.9

5.9

12.0

8.8

5.5

12.1

8.7

Caud. Ped. Depth

6.3

2.3

6.3

4.3

2.5

6.1

4.2

Snout

4.5

1.5

4.5

2.8

1.7

3.5

2.6

Eye

2.8

1.5

2.8

2.3

1.6

3.2

2.2

Jaw

5.6

1.7

5.9

3.4

1.7

4.3

2.8

Interorbit

4.6

1.4

4.7

3.0

1.6

4.0

2.7

Pectoral

8.8

4.2

9.0

6.7

4.4

8.9

6.5

Pelvic

6.4

3.0

6.6

4.7

2.9

6.4

4.6

Caudal

10.9

5.6

11.0

8.5

5.8

10.8

8.2

Longest D1 spine

5.9

2.8

5.9

3.7

2.6

4.6

3.6

H.K. Larson

base. Inner edge of shoulder girdle smooth with
no bony ridge or fleshy knobs or flaps. Gill
rakers on outer face of first arch very short
unspined knobs, longest raker by angle of arch;
outer rakers on other arches smaller than those
on first arch, outer rakers on fourth arch rudi¬
mentary; inner rakers on other arches nearly
twice the length of first arch inner rakers. Tongue
large, round to bluntly rounded. Teeth in female
slightly smaller than those of male. Teeth in
outermost row in upper jaw usually larger than
others, stout and curved or almost upright; be¬
hind this row, two to three rows of slightly
smaller stout curved teeth; rows narrow toone or
two at side of jaw. Teeth in lower jaw in four to
five rows; teeth arranged as in upper jaw apart
from outermost row being not much larger than
those in rows behind it.

Predorsal scales small, cycloid, usually reach¬
ing forward to above preopercular margin, often
anteriormost rows of scales placed irregularly,
somewhat scattered; predorsal scales in holotype
do not quite reach to above preopercular edge
(two specimens have predorsal midline com¬
pletely naked). Operculum naked or with patch
of small cycloid scales on upper third to half
(sometimes only one or two scales present).
Cheek always naked. Pectoral base with few
cycloid scales, occasionally naked. Prepelvic
area with small cycloid scales posteriorly (ante¬
rior half of breast usually naked). Belly with
isolated patch of weakly ctenoid scales under
pelvics; remainder cycloid. Ctenoid scales on
side of body in narrow wedge extending forward
to behind pectoral fin, usually broken into patch
of ctenoid scales behind pectoral fin and weakly
ctenoid area on caudal peduncle only (as in
holotype).

Head pores absent as in all Chlamydogobius.

Sensory papillae pattern longitudinal, as in Fig.
8. Rear portion of cheek row c indistinguishable
from anterior part of row b (possibly absent). Two
or three stows on snout; rows close togetherand
composed only of one papilla each. Mental row
/consists of two or three papillae only.

Coloration of fresh material. From colour
slides of two fish (dorsal views only) taken in
field conditions. Photographed specimens im¬
mature; do not show any bright blue or yellow
colouring in dorsal fin.

Background colour of head and body light
yellowish brown, with six brown to greyish
brown square saddles across back and one across
nape and opercles; saddles mostly composed of

Fig. 8. Chlamydogobius japalpa n. sp., papillae pattern.
Scale bar - 1 mm.

blotches and dark vermiculations; anterior part
of head with vermiculate greyish brown mar¬
bling and small spots. Dorsal saddles partly
connected to each other by irregular lines and
small blotches. Anterior half of body generally
darker than posterior half. Dorsal saddle across
nape mostly broken up into darker brown
vermiculations. Anterior half of head relatively
unmarked, with most distinct pattern being dark
brown vermiculations on snout and anterior part
of interorbital; dark brown bar extends from
front of eye to upper lip; part of dorsal snout
vermiculation forms intense dark irregular line
to snout tip. Fins pale brownish to fawn; axil of
pectoral base pale with dark brown
vermiculations.

Coloration of preserved material. Preserved
colour not greatly different from living speci¬
mens. Background colour yellowish to yellow¬
ish brown, with darker brown complex mottling
and irregular spots and broad streaks. Seven
brown dorsal saddles present, sometimes diffi¬
cult to distinguish, often broken up into series of
vermiculate blotches and irregular blotchy lines
(saddles more easily distinguished in dorsal
view). Along mid-side of body, series of six or
seven rectangular brown blotches, blotches of¬
ten interconnected above and below by irregular
lines. Pectoral base pale yellowish brown with
upper half of base marbled or blotched. At cau¬
dal base, but not extending onto fin, distinct
blackish brown vertical bar or Y-shaped blotch
present. Anteriormost dorsal saddle, which
crosses nape and onto opercle, usually com¬
posed of dark brown vermiculations and small
marbled blotches; pale, relatively unpatterned
area usually present on top of head from anterior
to preopercular margin to rear of interorbital

A review of the Australian endemic gobiid genus Chlamydogobius

space. Interorbital and top of snout covered with
dark brown complex vermiculate lines and
blotches; usually dark line from mid-snout to
snout tip, dorsal to posterior nostril. Broad brown
bar from front lower edge of eye to upper lip;
diffuse brown bar from ventral edge of eye
across cheek to fade out behind rictus; some¬
times another bar extends from ventral rear edge
of eye back to ventral edge of preopercle. Rear
half of preopercle variably blotched with brown
patches, sometimes pale. Underside of head and
body pale whitish yellow. Entire lining of body
cavity dense black.

Fins darker and more intensely marked in
males than in females. First dorsal fin greyish to
brownish grey, with sub-marginal whitish band
(probably white or yellow in live mature males);
below whitish band, broad dark grey to black
band, most intense posteriorly (probably blue in
live mature males). In mature males, second
dorsal and anal fins dusky greyish with con¬
spicuous white margin; in females and immature
males, second dorsal and anal fins translucent
whitish to translucent, fin rays brownish, with
dusky brownish blotches usually variably present
along fin bases. In mature males, caudal fin plain
dusky grey with narrow whitish margin. In fe¬
males and immature males, caudal fin translu¬
cent with many rows of small, vertically aligned,
diffuse brown spots; spots sometimes fused to-
gethertoform irregularlines. Pectoral fins trans¬
lucent to dusky. Pelvic fins whitish, or dusky
with whitish margin and frenum.

Comparisons. See under C. eremius (the most
similar species) for comparisons. In C. japalpa
the head markings are more fine and vermiculate
than in C. eremius , which often consist of mar¬
bled blotches.

Distribution. Found only in the Finke River
system of the Northern Territory, from Ormiston
Gorge to Finke River National Park.

Ecology. This species has been collected in
shallow pools with rock, sand or fine gravel
substrate; the fish were hiding among detritus/
mulm accumulated over sand. Fish in Ormiston
Creek were observed to be active only at night,
possibly to escape predators or high daytime
temperatures (or both). A photograph of typical
habitat is on the front coverof Larson and Martin
(1990); the holotype and other specimens from
NTM S.l 1436 are from the site illustrated.

Remarks. This species is considered to be
probably the same species as C. eremius by
Adams (pers. comm.) due to similar percentages
of fixed difference obtained by isozyme

electrophoresis. Morphologically, the two are
distinguishable and are here considered separate
species (see remarks under C. eremius).

Etymology. From the Western Aranda name
Japalpa , given to the part of the Finke River
which extends through what is now called Glen
Helen Gorge, as given by Strehlow (1947,1971).
The main waterhole in this gorge is just down¬
stream of the Ormiston Creek type locality, and
is a significant site in Aranda traditions. The
Finke is also “... said to be the oldest river on
Earth.” (Kotwicki 1989).

Chlamydogobius micropterus n. sp.

(Figs 3-4, 9-10, Tables 2-4, 7)

Chlamydogobius sp. nov. - Glover 1989: 98,

110 .

Chlamydogobius sp. A - Wager and Jackson
1993: 85.

Chlamydogobius n. sp. - Harris 1987: 8;
Jackson 1993: 23.

Chlamydogobius sp. - Morton etal. 1995:53,
119

Material examined. 37 specimens (9.5-23).
ELIZABETH SPRINGS, QUEENSLAND:
HOLOTYPE - QM 1.25096, 22.5 mm male,
Springvale Station, coll. J. Covacevich, P.
Couper, 27 April 1988. PARATYPES - AMS
1.25261-001, 3(8.5-20.5), Springvale, coll. W.
Ponder, P. Colman, 10 September 1984; AMS
1.25256-001,6(10.5-21.5), same data as preced¬
ing; QM 1.29552,27(9.5-23), same locality data
as holotype.

Diagnosis. A small Chlamydogobius with first
dorsal spines III-VI (mode V; pterygiophore
formula 4-2210 or 4-22000); second dorsal rays
1,6-8; anal rays 1,5-7; pectoral rays 11-13; longi¬
tudinal scales 35-47;TRB 12-17; predorsal scales
14-22, small, reaching to above preopercular
margin or further; scales on body mostly cycloid,
ctenoid scales restricted to patch under pectoral
fin; fins small, especially first dorsal; preserved
colour pale with indistinct broken-up rows of
light brown spots and mottling; known only
from Elizabeth Springs and associated pools on
Springvale Station, western Queensland.

Description. Based on 24 specimens, 12-23
mm SL. Counts of holotype indicated by asterisk
(Fig. 9).

First dorsal III* (1), IV (10), V (12), VI (1);
second dorsal 1,6-8 (mean 1,7*); analI,5-7 (mean
I,6*),pectoralrays 11-13 (mean 12*), segmented
caudal rays 15-17, mean 16*; caudal ray pattern
7/6 (4), 8/6 (4), 8/7* (11) or 9/7 (5); branched

H.K. Larson

Fig. 9. Holotype of Chlamydogobius micropterus n. sp., QM 1.25096, male.

caudalrays 13-16(mean 15*);longitudinal scale
count 35-47 (mean 41,43 in holotype); TRB 12-
17 (mean 15, 16 in holotype); predorsal scale
count 14-22* (mean 18); circumpeduncular scales
16-23 (mean 20,22 in holotype). Gill rakers on
outer face of first arch ranging from 1 +6 to 2+6
(mode). Pterygiophore formula 4-2210 (3), 4-
22000(2). Vertebrae 10+17(2), 10+18(7), 11+18
(3). No information available on the shape of
neural spines of first few vertebrae; X-rays not
sufficiently clear. One epural (9). One (1) or two
(6) anal pterygiophores before haemal spine of
first caudal vertebra.

Body somewhat rounded anteriorly, com¬
pressed posteriorly. Head rounded, wider than
deep, but not greatly so, HL 3.3-3.6 (mean 3.5)
in SL. Depth at posterior preopercular margin
1.1-1.7 (mean 1.6) in HL. Width at posterior
preopercular margin 1.3-1.5 (mean 1.4) in HL.

of about 25° with body axis; jaws reach to below
front half of eye in both sexes. Lips usually
smooth, fleshy fimbriae may be discernible; lower
lip free at sides, fused across front. Upper jaw
2.7-3.5 (mean 3.3) in HL. Eyes lateral, high on
head, top usually forming part of dorsal profile,
3.2-4.2 (mean 3.7) in HL. Snout rounded to
somewhat flattened, not overhanging upper lip,
3. l-4.4(mean 3.8) in HL. Interorbital broad, flat,
2.9-3.8 (mean 3.3) in HL. Top of head above
preopercular margin up to snout with very fine
villi sometimes present. Body depth at anal ori¬
gin 4.6-6.3 (mean 5.1) in SL. Caudal peduncle
compressed, length 3.1-3.7 (mean 3.3) in SL.
Caudal peduncle depth 6.3-8.1 (mean 7.1) in SL.

First dorsal fin reduced, somewhat pointed,
longest spine not much longer than eye width;
second or th ird spines longest or subequal; spines
no different between males and females; spines

Mouth terminal, slightly oblique, forming angle fall far short of second dorsal fin origin when

Table 7. Measurements (mm) of Chlamydogobius micropterus n. sp.

Character

Holotype

Minimum

Males

Maximum

Mean

Maximum

Females

Minimum

Mean

Head Length

6.8

3.8

6.8

5.3

3.7

6.7

4.7

Head Depth

4.2

2.6

4.3

3.4

2.6

4.0

3.1

Head Width

4.8

2.8

5.0

3.9

2.8

4.7

3.4

Body Depth

4.8

2.3

4.8

3.6

2.4

4.9

3.2

Body Width

3.2

1.5

3.5

2.5

1.6

3.5

2.2

Caud. Ped. Leng.

6.1

3.9

6.9

5.4

3.8

7.2

4.9

Caud. Ped. Depth

3.3

1.8

3.3

2.6

1.8

3.4

2.3

Snout

2.0

0.9

2.0

1.5

0.9

1.9

1.2

Eye

1.8

1.1

1.8

1.5

1.1

1.6

1.3

Jaw

2.3

1.1

2.5

1.7

1.1

1.2

1.5

Interorbit

2.2

1.0

2.2

1.6

1.1

2.2

1.5

Pectoral

4.7

2.6

4.7

3.6

2.7

5.0

3.4

Pelvic

3.2

1.7

3.2

2.4

1.7

3.3

2.2

Caudal

5.9

3.7

5.9

4.8

3.5

6.7

4.5

Longest D1 spine

1.9

1.0

1.9

1.5

1.4

1.9

1.5

A review of the Australian endemic gobiid genus Chlamydogobius

depressed. First dorsal spine always shorter than
next two. Second dorsal spine length 10.4-14.5
(mean 12.3) in SL. Third dorsal spine length
11.8-14.5 (mean 12.8) in SL. Second dorsal and
anal Fms very low, short-based, posteriormost
rays longest, rays fall far short of caudal fin base
when depressed; length of second dorsal fin fits
into space between tip of last fin ray and caudal
base. Pectoral fin rounded, central rays longest,

4.1 -5.6 (mean 5.0) i n SL; rays usu ally al 1 branched
(uppermost ray may be unbranched). Pelvic fins
short, rounded to oval, may reach half to two-
thirds of distance to anus, 6.5-8.8 (mean 7.6) in
SL. Caudal fin rounded, 3.4-4.6 (mean 3.7) in
SL.

No mental frenum, chin smooth. Anterior
nostril in very short tube placed on preorbital
edge, tube oriented down and forward, preorbital
may be produced forward slightly to accommo¬
date nostril. Posteriornostril small,round, placed
between front margin of eye and preorbital,
nostril closer to eye than preorbital edge. Gill
opening restricted to pectoral base. Inneredge of
shoulder girdle smooth with no bony ridge or
fleshy knobs. Gill rakers on outer face of first
arch small fleshy unspined knobs, longest raker
near angle of arch; rakers on inner face of first
arch small but broader than those on outer face;
inner rakers on other arches nearly twice length
of first arch inner rakers; outer face rakers of
fourth gill arch rudimentary to absent. Tongue
usually blunt. Outer teeth in upper jaw largest,
pointed, curved or almost straight, behind this
row are about two rows of small sharp teeth;
rows narrow to one or two at side of jaw (no
sexual dimorphism); tips of teeth (especially
outer row) often tinted translucent pale brown.
Lower jaw with about four rows of small pointed
teeth across front, outermost row oriented nearly
upright with tips turned inward; usually only one
row of teeth at side of jaw; tips of teeth often
tinted translucent pale brown.

Predorsal scales small, cycloid, evenly sized,
usually reaching forward to at least above
preopercular margin or to behind eyes.
Operculum with patch of small cycloid scales on
upper third to half. Cheek always naked. Pecto¬
ral base covered with small cycloid scales.
Prepelvic area at least with small cycloid scales
posteriorly, anterior portion often naked. Belly
with isolated patch underpelvicsusually ctenoid
(in 10), rest cycloid; or belly scales all cycloid
(10). Ctenoid scales on side of body restricted to
small patch beneath pectoral fin; ctenii weak.

Fig. 10. Chlamydogobius micropterus n. sp., papillae pat¬
tern. Scale bar - 1 mm.

Head pores absent as in all Chlamydogobius.

Sensory papillae pattern longitudinal, as in
Fig. 10. Cheek rows b and c reduced, broken.
Two j rows on snout, consisting of one or two
papillae each. Mental / row of one papilla on
each side.

Coloration of fresh material. From colour
slide of captive male specimen.

Head and body pale greyish yellow, with
white abdomen. Rear portion of some scales
yellowish white, forming irregular marbled pat¬
tern over upper half of body. Dense, very dark
brown spots widely scattered over body, few on
dorsal surface of head (apparently restricted to
scaled portion of nape). Brown spots larger and
more evenly spaced along posterior mid-side of
body; largest spot at centre of caudal fin base.
Head mostly plain, with broad dark-margined
pale brownish bar extending from below eye to
end behind jaws. Opercle with brownish mark¬
ings and yellowish white-margined scales
present. Snout, jaws and lower half of head pale
yellowish white. Iris pale golden with brownish
margin.

First dorsal fin (very small) appears to be
yellowish grey anteriorly with large bright sky
blue spot occupying most of fin (fin partly folded).
Seconddorsal fin with membranes mostly trans¬
lucent, some dusky barring visible; fin rays yel¬
lowish grey. Anal fin grey with white margin (fin
mostly obscured by body). Caudal fin translu¬
cent yellowish grey, uppermost fin rays nearly
whitish. Pectoral fin folded, but appears whitish.
Pelvics not visible.

Coloration of preserved material. All avail¬
able preserved specimens pale; no males in breed¬
ing colouring present.

Head and body pale yellowish white to pale
brownish yellow, with indistinct light brown

H.K. Larson

markings; peritoneum shows through body wall
as silvery white or shiny purplish brown, de¬
pending on state of preservation. Upper half of
body light dusky, with irregular and very vari¬
able light brown mottling and speckling, scale
margins often outlined with light brown, form¬
ing irregular zig-zag streaks. Five to seven vari¬
able brown blotches or short bars may cross
dorsal mid-line from first dorsal fin origin back¬
ward; nape generally plain. Mid-line of body
sometimes shows diffuse series of light brown
spots or short vertical dashes; most distinct mark
is roughly rounded light brown spot at centre of
caudal base, spot may be partly joined to two
diffuse brownish spots on base of fin itself.
Pectoral base dusky. Top and sides of head
dusky pale brownish with slightly darker brown
indistinct mottling; usually three markings dis¬
tinguishable: brown area from front of eye for¬
ward to preorbital edge below anterior nostril,
brown blotch from ventral edge of eye reaching
down to behind rictus and diffuse brown blotch
from rear edge of eye across preopercle. Lips
pale to dusky, chin pale. Peritoneum black inter¬
nally.

First dorsal fin pale to dusky with dark brown
to dark grey blotch posteriorly. Second dorsal fin
translucent to whitish with brownish fin rays.
Anal, pectoral and pelvic fins whitish. Caudal fin
translucent to dusky with indistinct brownish
barring orrowsof spots proximally, fin unspotted
distally; two brown spots on either side of fin
base, spots sometimes coalesced into broad dusky
brownish vertical bar.

Comparisons. This species is distinctive in
possessing a greatly reduced (reduced in size as
well as number of spines) first dorsal fin; only
one specimen had six spines. It also differs in
colour pattern, apparently being pale with dark
fine spots which do not form marbling or
vermiculation as in most of the other species.

Distribution. Specimens are only known from
spring-fed pools on the Springvale Station (Spring
Valley on older maps), western Queensland (23°
30' S 140° 35' E).

Ecology. There appears to be very little infor¬
mation about the environment this species in¬
habits, other than it being extremely vulnerable
to destruction. Wager and Jackson (1993) indi¬
cate that the water level and flow of Elizabeth
Springs “...is decreasing due to a reduction in the
water table caused by water extraction through
artificial bores”.

Remarks. Theholotype, although possessing
the least number of dorsal spines, was chosen

due to its good condition and fairly dark pigmen¬
tation (dark when compared with other speci¬
mens of this apparently pale species).

Wager and Jackson (1993) discuss the conser¬
vation status of this species and indicate that it
deserves high priority for action, before the wild
population becomes extinct due to a combina¬
tion of water draw-down by artificial bores and
habitat destruction by stock trampling; they pro¬
pose several management actions which should
be taken. Apparently Gambusia has not yet in¬
vaded this area.

To date, C. micropterus remains on the Aus¬
tralian Society for Fish Biology’s 1993 Threat¬
ened Fishes (sixth) Supplement list as “endan¬
gered”; oneof the 11 Australian freshwater fishes
(and the only goby) to be so honoured. It has
been on the ASFB’s Threatened Fish listsupple-
ments, under a variety of categories, since 1987
when it was listed as “potentially threatened”
(Harris 1987).

Etymology. From the Greek, micropterus
(small fin), in reference to the reduced first
dorsal fin.

Chlamydogobius ranunculus n. sp.

(Figs 3-4, 11 -12, Plate 2, Tables 2-4, 8)

Mugilogobius sp. 9 - Gee and Gee 1991: 19,
21-26.

Material examined. 167 specimens (4-31).
HOLOTYPE - NTM S. 11427-001, 27.5 mm
male, partly dried-up buffalo wallow, Beatrice
Lagoon, Adelaide Riverdrainage, 12°37’S 131°
21 ’E, coll. H. Larson, 20 May 1984.
PARATYPES - QUEENSLAND: AMS 1.22959-
001, 5(22-25.5), lagoon behind wharf,
Townsville, coll. J. Gee, 16 June 1981; QM
1.19003, 1(31), Norman River near Karumba,
coll. D.J. Russell, 11 June 1981; QM 1.19005,
1(27), same data as preceding; AMS 1.20928-
001,27(9.5-17.5), Smith Point, Prince of Wales
Island, Torres Strait, coll. D.F. Hoese, 1979.
NORTHERN TERRITORY: NTM S. 11427-002,
7( 19-27.5), same locality data as holotype; NTM
S.11509-007,118(4-31), bombholes at Leanyer
Swamp, coll. NT. Fisheries, 9 October 1984;
AMS 1.32051-022, 5(20.5-31), Alligator River
mouth, coll. T. Davis, April 1979; ex AMS
1.32051 -020,2(23-27), cleared and stained. Al¬
ligator River, coll. T. Davis, April 1979.

Other material examined (but not used in
description). 150 specimens from the following
localities. AMS 1.26677-007, 14, Hinchinbrook
Channel, Queensland; NTM S.11935-001, 1,

A review of the Australian endemic gobiid genus Chlamydogobius

Fig. 11. Holotype of Chlamydogobius ranunculus n. sp., NTM S.l 1427-001, male.

McArthur River, NT; NTM S.l 1510-003, 3,
Leanyer Swamp, NT; NTM S.l 1509-006, 61,
Leanyer Swamp, NT; AMS 1.19229-001, 1,
Marjarie Creek, NT; AMS 1.32051-012,29, Al¬
ligator River drainage; AMS 1.32051-027, 5,
Alligator Riverdrainage; AMS 1.32051-020,12,
Alligator Riverdrainage; AMS 1.32051-019,10,
Alligator River drainage; AMS 1.32051-011,4,
Alligator River drainage.

Diagnosis. A moderately sized
Chlamydogobius with second dorsal rays 1,6-
8mean; anal rays I; pectoral rays 11-13; longitu¬
dinal scales 32-52; TRB 12-17; predorsal scales
small, reaching to above preopercular margin
and or further forward; ctenoid scales on body
usually consist of narrow wedge along mid-side
or as two discrete patches; head rounded, almost
as deep as wide; usually 27 vertebrae; estuarine,
known from low salinity muddy habitatsin north¬
ern Australia (Queensland and the Northern Ter¬
ritory).

Description. Based on 54 specimens, 14-31
mm SL. Counts of holotype indicated by asterisk
(Fig. 11).

First dorsal V (1), VI* (48); second dorsal 1,6-
8 (,5-71,6*); anal 1,5-7 (mean 1,6*), pectoral rays
1 l*-13(mean 12), segmented caudal rays 15-17
(mean 16,15 in holotype); caudal ray pattern 7/
6* (11), 7/7 (4), 8/6 (11), 8/7 (22) or 9/7 (1);
branched caudal rays 11-16 (mean 14, 13 in
holotype); unsegmented (procurrent) caudal rays
6/6 (2), 6/7 (1), 7/6 (1); longitudinal scale count
32-52 (mean 42, 39 in holotype); TRB 12-17
(mean 14,13 in holotype); predorsal scale count
3-20 (mean 14, 16 in holotype, 0 in two speci¬
mens); circumpeduncular scales 15-23 (mean
20, 18 in holotype). Gill rakers on outer face of

first arch ranging from 1+6 to 4+7 (mode 2+8).
Pterygiophore formula 3-12210 (15), 3-12200
(1), 3-122110 (1). Vertebrae 10+16 (5), 10+17
(6), 11+16 (12), 11 + 17 (4), 11+18 (1). Neural
spines of first three vertebrae split or expanded at
tip (11) or narrow and pointed (2). One (18) or
two (3) epurals. One (11) or two (13) anal
pterygiophores before haemal spine of first cau¬
dal vertebra.

Body compressed (less so anteriorly). Head
rounded, usually wider than deep, HL 3.0-3.6
(mean 3.4) in SL; sometimes almost square in
cross-section. Depth at posterior preopercular
margin 1.4-1.8 (mean 1.6) in HL. Width at pos¬
terior preopercular margin 1.2-1.6 (mean 1.4) in
HL. Mouth terminal to subterminal, somewhat
oblique, forming angle of about 25° with body
axis; jaws generally reach to below mid-eye in
both sexes; in smaller females, jaws may reach to
below front half of eye. Lips less fleshy than in
other Chlamydogobius, usually smooth, fleshy
fimbriae may be present mostly on inner edges of
upper I ip, less often on lower lip (present on both
lips in holotype); lower lip free at sides only,
broadly fused across front. Upper jaw slightly
longer in males than in females, 2.0-3.5 (mean
2.9 in females, 2.4 in males) in HL. Eyes lateral,
high on head, top may form part of dorsal profile,
2.8-4.6 (mean 3.7) in HL. Snout somewhat steep,
rounded, 2.9-4.1 (mean 3.6) in HL, occasionally
somewhat inflated and may partially overhang
upper lip. Interorbital broad, flat, 2.9-5.4 (mean
3.9) in HL; in large specimens, eyes may be
placed high on head with flesh surrounding eye
giving interorbital concave appearance. Top of
head, from above preopercular margin up to
snout, may have very fine dark-pigmented villi

H.K. Larson

present. Body depth at anal origin 4.3-8.8 (mean
5.1) in SL. Caudal peduncle compressed, length

3.2- 7.5 (mean 3.9) in SL. Caudal peduncle depth

6.3- 12.1 (mean 7.4) in SL.

First dorsal fin low, rounded, third and fourth
spines longest or subequal; spines often slightly
longer in males than females; in mature males,
spines barely reach second dorsal origin when
depressed; fin usually falls short of second dor¬
sal in females. First dorsal spine always shorter
than next three. Second dorsal spine length 9.8-
10.4(mean 10.1) in SL. Third dorsal spine length
7.6-10.2 (mean 8.9) in SL. Fourth dorsal spine
length 7.2-10.8 (mean 9.0) in SL. Second dorsal
and anal fins low, posteriormost rays longest,
rays fall well short of caudal fin base when
depressed, in both sexes. Pectoral fin rounded,
central rays longest, 3.8-5.1 (mean 4.4) in SL;
rays usually all branched. Pelvic fins short, usu¬
ally oval, may reach half distance to anus, 5.4-
7.9 (mean 6.5) in SL. Caudal fin broad, rounded,
2.9-3.9 (mean 3.5) in SL.

No mental frenum, chin smooth. Anterior
nostril in very short tube, placed on preorbital
edge just behind upper lip, tube oriented down
and forward, preorbital curved forward to ac¬
commodate nostril. Posterior nostril small,
rounded to oval, placed halfway between front
margin of eye and preorbital. Gill opening re¬
stricted to pectoral base. Inner edge of shoulder
girdle smooth with no bony ridge or fleshy
knobs. Gill rakers on outer face of first arch very
short unspined knobs, longest raker near angle of
arch, sometimes rudimentary rakers alternate
with “full-sized” rakers on lower limb, outer
rakers on fourth arch very rudimentary but vis¬
ible; rakers on inner face of first arch also stubby;

inner rakers on other arches unspined, short, bht
nearly twice length of first arch inner rakers.
Tongue broad, usually concave (nearly bilobed
in several specimens) to bluntly rounded. Outer
teeth in upper jaw largest, curved, with bluntly
pointed tips; behind this row are two or three
rows of smaller teeth; rows narrow to one or two
at side of jaw. Lower jaw with two or three rows
of smaller pointed teeth across front, outermost
row oriented upright, teeth of inner rows tend to
curve inward; usually only one or two rows Of
teeth at side of jaw (no sexual dimorphism). Tips
of teeth (especially outermost) in both jaws often
tinted translucent orange or honey brown.

Predorsal scales cycloid, small, evenly sized,
usually reaching forward to above preopercular
margin or slightly further, predorsal scales in
holotype just reach over preopercular edge.
Operculum with small cycloid scales on upper
third to half; sometimes only few scales present.
Cheek always naked. Pectoral base covered with
cycloid scales. Prepelvic areanaked, orwith few
cycloid scales before pelvics, or occasionally
completely covered with scales. Belly usually
with isolated patch under pelvics of weakly
ctenoid scales, rest cycloid; some specimens
with entire belly cycloid. Ctenoid scales on side
of body in narrow wedge along midline, or more
usually, broken into patch of scales behind pec¬
toral fin, with remaining ctenoid scales begin¬
ning below second dorsal fin origin or further
back on caudal peduncle (ctenii weak, often
obscured by skin and mucous coat).

Head pores absent as in all Chlamydogobius.

Sensory papillae pattern longitudinal, as in
Fig. 12. Cheek rows b and cp short and/orbroken
into short sections; row c usually only consists of

Tabic 8. Measurements (mm) of Chlamydogobius ranunculus n. sp.

Character

Holotype

Minimum

Males

Maximum

Mean

Maximum

Females

Minimum

Mean

Head Length

8.3

4.1

9.6

7.4

5.1

9.1

6.9

Head Depth

5.6

2.5

6.0

4.7

3.2

6.4

4.5

Head Width

6.4

2.8

7.0

5.4

3.5

7.7

5.2

Body Depth

5.7

2.9

6.8

5.0

3.6

7.2

5.0

Body Width

3.6

1.6

3.6

2.8

2.0

3.5

2.8

Caud. Ped. Leng.

7.6

4.1

8.1

6.6

4.7

8.0

6.5

Caud. Ped. Depth

4.4

2.0

4.5

3.5

2.5

5.0

3.4

Snout

2.5

1.0

2.9

2.1

1.3

2.9

1.9

Eye

2.2

1.2

2.4

2.0

1.4

2.5

1.9

Jaw

3.5

1.6

4.7

3.1

1.5

4.1

2.5

Interorbit

1.9

0.9

2.9

2.0

1.2

3.0

1.8

Pectoral

6.4

3.5

7.7

5.9

4.1

7.1

5.4

Pelvic

4.8

2.6

5.1

4.1

2.8

5.3

3.7

Caudal

8.1

4.3

9.3

7.3

5.4

8.7

6.9

Longest D1 spine

3.6

2.6

4.0

3.1

2.4

3.5

2.7

A review of the Australian endemic gobiid genus Chlamydogobius

Fig. 12. Chlamydogobius ranunculus n. sp., papillae pat¬
tern. Scale bar - 1 mm.

one papilla. Two or three s rows on snout, rows
close together, usually consisting of one papilla
each; anteriormost row often consists of two
papillae. Mental/row with two or four papillae.

Coloration of fresh material. From slides of
freshly dead specimens (Plate 2), and one living
specimen (Plate 2).

Mature male. Head and body brownish grey,
pale yellowish white to whitish grey on under¬
side of head and abdomen. Six or seven brown¬
ish grey (darker than background colour, but not
greatly so) bars or saddles across back to mid¬
side of body; posteriormost bars somewhat ob¬
lique. Indistinct brownish grey spot at centre of
caudal base. Head relatively plain brownish grey
with no dark spots or bars other than two diffuse
dark bars from eye extending to upper lip and to
behind rictus respectively; lower half of opercle
very pale bluish grey. Iris marbled with dark
brown and gold. Pectoral base mostly brownish
grey, whitish yellow anteriorly.

First dorsal fin dark grey along outer margin
and on lower half of fin. Broad maize yellow
submarginal band, below this and to rear of fin,
light blue to greyish blue oval spot, spot edged
with black ventrally; fin rays dark grey to black.
Second dorsal fin dark grey to blackish with
broad white margin, fin rays grey; lower half of
fin with broad band of greyish blue, blue brighter
toward rear half of fin; on fin membranes at
about middle of fin, series of vertically oriented
black blotches present. Anal fin similarly col¬
oured to second dorsal fin but with no series of
black blotches across middle of fin; greyish blue
on proximal half of fin brighter than in second
dorsal fin. Caudal fin grey with indistinct, verti¬
cally oriented oval dark grey blotch on each side
of fin base; vertically oriented rows of small dark
spots and blotches extend half length of fin,

growing more indistinct distally; narrow dull
white margin present. Pectoral fin membranes
translucent, fin rays greyish to yellowish white.
Pelvic fins yellowish white.

Female or immature male. Head and body
translucent yellowish grey, underside of head
and abdomen whitish. Body with indistinct grey
to darker yellowish grey saddles and mottling;
diffuse greenish grey streak extends from behind
eye along upper half of body to below origin of
second dorsal fin; scattered scales along mid¬
side of body with bluish white spots. Dorsal
surface and upper sides of head with greenish
grey spots, small blotches and vermiculate lines;
lower half of opercle greenish silvery white. Iris
pale gold with narrow brown ring around pupil.
Peritoneum (through body wall) with upper half
pinkish white overlain by diffuse blackish nar¬
row bars, ventral half whitish. Pectoral base
pinkish white, greyish close to ray bases.

First dorsal fin with narrow grey margin, broad
greyish yellow band across distal half of fin;
below this, bright pale blue rounded spot be¬
tween fourth and sixth spines; lower half of fin
translucent dusky grey. Second dorsal fin trans¬
lucent pale grey with pinkish white margin,
lowerhalfof fin irregularly marked with slightly
darker grey ish orpinkish white vermiculate 1 ines
or diffuse blotches. Anal fin dark grey with
bright white margin, proximal half of fin with
greyish white markings (form indiscernible from
photograph). Caudal fin translucent light grey
with white margin; most of fin with vertically
aligned rows of small grey or wh ite spots; diffuse
greenish grey dark spot on either side of fin base.
Pectoral fin membranes translucent; rays pale
translucent grey. Pelvic fins dark grey.

Coloration of preserved material. Males in
breeding colour very similar to pattern as de¬
scribed for live material. Head and body pale
yellowish brown to yellowish white (depending
on preservation), with brown to light brown
markings; spots and bars more conspicuous in
preserved than live material. Pattern on body
variable and often difficult to distinguish, but
roughly formed by seven variably shaped (often
square to rectangular) brown blotches on upper
half of body and seven slightly darker brown
spots or elongate blotches along mid-side of
body (these two series of markings slightly off¬
set). Body markings sometimes partly joined by
diffuse lines and series of vague spots, forming
diffuse streaks; markings may be formed by
brownspots in scale centresor by brown margins

H.K. Larson

to scales. Predorsal and top of snout generally
plain brown, as is top of opercle; lower half of
opercle light brown; preopercle diffusely mot¬
tled or spotted with brown. Brown bar extends
from front of eye to upper lip (bar runs just below
nostrils so they appear to be in pale stripe be¬
tween dark top of snout and dark eye-bar); sec¬
ond brown bar extends from ventral edge eye to
just behind rictus; third indistinct bar or blotch
may extend from lower rear edge of eye across
cheek. Lower lip and chin speckled with brown;
underside of head and abdomen yellowish white
to yellowish brown. Pectoral base brown, or
brown on upper half and pale on lower. Caudal
base with brown spot at centre; spot not always
darker than other body markings. Internally,
peritoneum dark brown to black dorsally, fading
ventrally and on lower sides to light brown or
dusky grey.

First dorsal fin with greyish brown margin,
broad submarginal transparent to translucent
whitish band below this; lowerhalf of fin greyish
brown; usually dark grey to black spot on rear
half of fin. Second dorsal and anal fins light
greyish brown with broad translucent margins;
second dorsal (less often, anal) fin may have one
or two rows of darker brownish vertically elon¬
gate blotches on membrane; blotches in series in
middle of fin. Caudal fin dusky grey with rows of
vertically aligned diffuse small dark spots, and
narrow whitish or translucent margin; some¬
times spots nearest caudal base dark brown.
Pectoral fins translucent to yellowish white, rays
dusky. Pelvic fins translucent to yellowish white,
bases of rays dusky.

Comparisons. This species differs from other
Chlamydogobius by possessing an equal number
of soft dorsal and anal fin rays, in having the
lowest pectoral ray count (11-12), and being the
only coastal species of the genus.

Distribution. Specimens are so far known
only from northern Queensland and the North¬
ern Territory, Australia.

Ecology. Northern Territory specimens have
been collected from shallow, muddy, low salin¬
ity (0-9%o), coastal habitats. A number have
come from rather precarious habitats such as tiny
muddy creeklets draining mangrove samphire
plain or freshwater floodplain, artificial habitats
such as flooded bomb craters, water buffalo
wallows, and concrete town drains (in general,
the worse-looking the creek or puddle, the more
likely that C. ranunculus will be present). No
information has yet been obtained on this spe¬

cies’ thermal and salinity tolerances, but they are
likely to be wide, as may be the case with all the
freshwater species.

Gee and Gee (1991) describe methods of
aquatic surface respiration used by this species
(reported as Mugilogobius sp. 9) when it was
exposed to low oxygen levels; the fish used
bubble-holding, with the body held in an arched
or vertical position.

Neil Armstrong of the Australia New Guinea
Fishes Association successfully induced (by
abundance of food and TLC) captive specimens
to spawn (1994; pers. comm.). About 30 large
eggs were laid, which took about nine days to
hatch at 25°C; the male guarded the eggs. Shortly
after hatching, the young fish were observed
clinging to the walls of the aquarium and ap¬
peared to be feeding on the surface film.

Remarks. This species, although displaying
some specialisations (such as the even ratio of
dorsal to anal rays) is likely to be closer to the
marine ancestor of the group than any of the
freshwater Chlamydogobius. The question of
the desert gobies’ ancestors have intrigued me
and Miller (1987) for some time (did
Chlamydogobius come from a temperate south¬
ern Australian group similar to Mugilogobius
paludis or its ancestor, or from a northern tropi¬
cal group?); the question is discussed later in this
paper.

This species has been identified in some mu¬
seum collections as Mugilogobius sp. 9 (or DFH
sp. 9, in reference to Doug Hoese of AMS).

Etymology. From the Latin ranunculus , mean¬
ing tadpole, a resemblance to which this rather
frog-headed goby displayed to the author upon
first their encounter, at the edge of a drying-up
water buffalo wallow.

Chlamydogobius squamigenus n. sp.

(Figs 3 -4, 13-14, Tables 2-4, 9)

Chlamydogobius n. sp. - Jackson 1993: 23.

Material examined. 46 specimens (14-39).
QUEENSLAND: HOLOTYPE - SAM F.6595,
34 mm male, “Western” spring, approximately
2.4 km NE of Edgbaston Station Homestead,
coll. J. Glover, T. Sim and T. Scott, 7 May 1989.
PARATYPES - SAM F.7676, 8(21 -34.5), same
locality data as holotype; SAM F.6738, 1(34),
small mound artesian spring approximately 3.3
km SSE of Edgbaston Homestead, coll. W.
Zeidler, 4 May 1988; SAM F.7184, 35 (14-39),
“Crossmoor Flowing Bore”, Crossmoor Station,

A review of the Australian endemic gobiid genus Chlamydogobius

Fig. 13. Holotype of Chlamydogobius squamigenus n. sp., SAM F.6995, male.

on Longreach-Muttaburra Road, coll. T. Sim
and P. Num, 3 June 1993; SAM F.7206, 1(35),
Crossmoor Station, Bore Number 1652, south
arm of drain, coll. T. Sim and P. Num, 5 June
1993.

Diagnosis. A small Chlamydogobius with sec¬
ond dorsal rays 1,6-8; anal rays 1,5-7; pectoral
rays 13-14; longitudinal scales 33-45; TRB 14-
17; 28 vertebrae; predorsal scales small, reach¬
ing past preopercular margin nearly to behind
eyes; preopercle with small cycloid scales at
least in patch by lower rear comer or below eye,
ctenoid scales on side of body weak, in two
separate patches; species known only from
springs and pools on Edgbaston and Crossmoor
Stations, Thomson River drainage, central
Queensland.

Description. Based on 32 specimens, 21-39
mm SL. An asterisk indicates counts of the
holotype (Fig. 13).

First dorsal V (5), VI* (25), VII (2); second
dorsal 1,6-8 (mean 1,7*); anal 1,5-7* (mean 1,6),
pectoral rays 12-14 (mean 13, 14 in holotype),
segmented caudal rays 15-18 (mean 16*); cau¬
dal ray pattern usually 8/7 or 9/7*; branched
caudal rays 14-17 (mean 16*); unsegmented
(procurrent) caudal rays usually indiscernible in
X-rays, 7/5 (1), 7/6 (1), 7/7 (1); longitudinal
scale count 33-45 (mean 39, 40 in holotype);
TRB 14-17* (mean 16); predorsal scale count
15-22 (mean 19, 21 in holotype);
circumpeduncular scales 17-21 (mean 19,18 in
holotype). Gill rakers on outer face of first arch
ranging from 0+6* to 2+7 (mode 2+6).
Pterygiophore formula 3-12210 (10), 3-11210
(2). Vertebrae 10+18 (15), 11+17 (2). Neural
spines of first few vertebrae narrow, pointed (7).

One broad epural, sometimes partly split (14).
Two (13) or one (2) anal pterygiophores before
haemal spine of first caudal vertebra.

Body somewhat rounded anteriorly, com¬
pressed posteriorly. Head wider than deep, HL

2.2- 33 (mean 3.0) in SL; in small specimens,
head almost square in cross-section. Depth at
posteriorpreopercular margin 1.5-1.8(mean 1.7)
in HL. Width at posterior preopercular margin

1.3- 1.6 (mean 1.4) in HL. Mouth subterminal,
slightly oblique, forming angle of about 20-23°
with body axis; jaws generally reach to below
fronthalfof eye or to front margin of eye; in large
males jaw may reach to below mid or rear half of
eye. Lips usually smooth, fleshy fimbriae some¬
times present on inner edges of upper lip; lower
lip free at sides, broadly fused across front.
Upper jaw 2.2-3.2 (mean 3.0 in females, 2.7 in
males) in HL. Eyes lateral, high on head, top
usually forming part of dorsal profile, 3.5-53
(mean 4.6) in HL. Snout rounded, slightly flat¬
tened to steep, 3.1-4.1 (mean 3.6) in HL.
Interorbital moderate tobroad,flat,2.8-4.1 (mean
3.5) in HL. Top of head above preopercular
margin up to front of interorbital with very fine,
usually widely scattered villi. Body depth at anal
origin 3.3-5.3 (mean 4.8) in SL. Caudal peduncle
compressed, length 2.7-4.3 (mean 3.7) in SL.
Caudal peduncle depth 4.7-7.4 (mean 6.8) in SL.

First dorsal fin low, rounded, third or fourth
spines longest or subequal; spines about the
same length in males and females; in both sexes,
spines fall short of second dorsal origin when
depressed (even in mature males). First dorsal
spine always shorter than next three. Second
dorsal spine length 10.3-17.0 (mean 10.5) in SL.
Third dorsal spine length 6.7-10.8 (mean 9.7) in

H.K. Larson

Table 9. Measurements (mm) of Chlamydogobius squamigenus n. sp.

Character

Holotype

Minimum

Males

Maximum

Mean

Maximum

Females

Minimum

Mean

Head Length

10.4

7.0

12.7

9.4

6.8

11.7

9.5

Head Depth

7.0

4.0

8.0

5.7

4.0

7.2

5.7

Head Width

8.3

4.7

9.7

6.7

4.7

8.5

6.6

Body Depth

7.7

4.3

7.9

5.9

4.3

8.0

6.1

Body Width

4.6

2.5

4.8

3.6

2.7

5.2

3.7

Caud. Ped. Leng.

9.1

6.1

10.2

7.7

6.0

9.8

7.8

Caud. Ped. Depth

5.4

3.0

5.7

4.2

3.1

5.5

4.3

Snout

3.3

1.9

3.9

2.6

1.7

3.5

2.7

Eye

2.2

1.6

2.5

2.0

1.6

2.4

2.1

Jaw

3.9

2.5

5.9

3.5

2.1

4.1

3.2

Interorbit

3.5

1.9

4.4

2.8

1.8

4.0

2.7

Pectoral

6.8

4.7

8.3

6.2

4.4

7.7

6.2

Pelvic

5.0

3.5

6.8

4.8

3.3

6.0

4.8

Caudal

8.3

5.4

9.8

7.4

5.8

9.0

7.4

Longest D1 spine

3.2

2.5

4.6

3.2

3.0

4.4

3.2

SL. Fourth dorsal spine length 8.4-13.6 (mean
10.2) in SL. Second dorsal and anal fins short-
based, low, posteriormost rays longer in mature
males; rays generally equal in others; rays never
reach caudal fin when depressed. Pectoral fin
broad, rounded, central rays longest, 3.3-5.0
(mean 4.6) in SL; rays usually all branched.
Pelvic fins short, oval, may reach half distance to
anus, 4.1-6.8 (mean 6.0) in SL. Caudal fin rec¬
tangular to rounded, 2.9-4.6 (mean 3.9) in SL.

No mental frenum, chin smooth. Anterior
nostril in short tube on preorbital edge just be¬
hind upper lip, tube oriented down and forward,
preorbital curved forward to accommodate nos¬
tril. Posterior nostril small, round, placed half¬
way between front edge of eye and preorbital.
Gill opening restricted to pectoral base. Inner
edge of shoulder girdle smooth with no bony
ridge or fleshy knobs or flaps. Gill rakers on
outer face of first arch low fleshy knobs, without
spines; longest raker near angle of arch; rakers
on inner face of first arch also short and stubby;
inner rakers on other arches nearly twice length
of first arch outer rakers. Tongue usually blunt or
rounded. Outer teeth in upper jaw largest, rela¬
tively slender and very slightly curved, behind
this row are three rows of small sharp curved
teeth; rows narrow to one or two at side of jaw
(no sexual dimorphism); tips of teeth sometimes
slightly tinted translucent honey brown. Lower
jaw with three or four rows of moderate sized,
curved pointed teeth across front, outermost row
largestand leastcurved; teeth of innerrows more
curved, tending to point posteriorly; usually only
one or two rows of teeth at side of jaw; tips of
teeth sometimes slightly tinted translucent honey
brown.

Predorsal scales cycloid, small, evenly sized,
usually reaching forward to past preopercular
margin, sometimes extending up to behind eyes.
Operculum with small cycloid scales on upper
third to half. Cheek at least with one or two
cycloid scales below eye or on rear part of
preopercle, usually distinct patch of scales
present; cheek apparently without scales in one
21 mm male specimen. Pectoral base covered
with cycloid scales. Prepelvic area covered with
small cycloid scales. Isolated area of ctenoid
scales covered by pelvic fins, rest cycloid. Body
scales mostly cycloid, with small separate patch
of ctenoid scales behind pectoral fin; weakly
ctenoid scales scattered along mid-line of caudal
peduncle or sometimes further; ctenoid scales
extend forward up to mid-body, at most.

Head pores absent as in all Chlamydogobius.

Sensory papillae pattern longitudinal, as in
Fig. 14. Cheek row c broken into two widely
separated portions; anterior part (below front

Fig. 14. Chlamydogobius squamigenus , papillae pattern.
Scale bar - 1 mm.

A review of the Australian endemic gobiid genus Chlamydogobius

edge of eye) of two or three papillae and poste¬
rior part of one or two papillae very close to, and
just below, anterior portion of row b. Two s rows
on snout; each row of one papilla each. Mental/
row consists of two papillae.

Coloration of fresh material.

Mature male. From colour slide. Head and
body dusky greyish yellow; head whitish yellow
underneath; peritoneum, through body wall, sil¬
very white. Indistinct narrow grey bars and
mottling across upper half of body. Head plain
greyish yellow; lips grey to whitish grey; iris
pale gold.

Dorsal fin blackish with narrow chrome yel¬
low submarginal band, band widest anteriorly,
becoming narrower and darker posteriorly; broad
vivid blue band across centre of fin; lower third
of fin blackish. Second dorsal and anal fins
blackish with broad bright greyish blue margin;
second dorsal fin with diffuse greyish blue band
across proximal half of fin. Caudal fin very dark
grey with narrow dull whitish grey margin; in¬
distinct vertically aligned rows of yellowish to
light grey spots present.

Coloration of preserved material. Head and
body dull yellowish white to greyish white with
brownish grey or grey scale margins, diffuse
narrow blotches and spots on upper half; lower
half of body, and underside of head, plain. Scales
on body diffusely outlined with grey or brownish
grey, forming indistinct bars, blotches and cross-
hatching which vary among specimens; from
dorsal view about six indistinct blotches or short
bars cross dorsal mid-line. Mid-side of body
with indistinct row of dark spots, only defined
spot in series is at mid-base of caudal fin. Pecto¬
ral base mostly pale, dusky patch across upper
part. Nape behind eyes, interorbital and top of
snout often with short, diffuse grey vermiculate
linesand spots. Broad diffuse brownish bar from
front of eye to upper lip, ending just below
anterior nostril tube. Second diffuse brownish
mark runs along lower edge of eye then curves
down to behind rictus.

First dorsal fin colour variable, usually trans¬
lucent to light grey with narrow dusky to black¬
ish margin, and blackish streak across middle to
upper half of fin, streak intensified as dense
black blotch posteriorly. Second dorsal and anal
fin whitish. Second dorsal may have series of
vertical dusky blotches forming band below
margin; other diffuse dusky grey blotches scat¬
tered irregularly over proximal half of fin. Cau¬
dal fin translucent to whitish, with many rows of

small vertically aligned dark grey or diffuse grey
spots; margin clear orwhite. Pectoral fin translu¬
cent whitish, rays dusky proximally. Pelvic fins
whitish; few heavily pigmented specimens have
dusky markings along rays but not frenum.

Mature male (SAM F.7206) similar to live
colour pattern described above, but dusky irregu¬
lar grey barring on upper sides more prominent.

Comparisons. Chlamydogobius squamigenus
is distinctive in having at least one or two cycloid
scales on the cheek (usually a patch of scales
present), unlike other species of the genus.

Distribution. Specimens are known only from
springs and pools at Edgbaston Station, NE of
Aramac, Queensland, and from bores and drains
on Crossmoor Station just NNE of Longreach;
both on the Thomson River drainage.

Edgbaston Station is at 22° 44' S 145° 25' E,
according to Ivantsoff et al. (1991). Although
there are many spring-fed pools on the station,
the number which contain C. squamigenus has
not been ascertained. “Crossmoor Flowing Bore”
from which mostofthe Crossmoor Station speci¬
mens came from, is at 22° 54' S 144° 35' E.

Ecology. This species was first collected from
a small artesian mound spring (at 22° 45' 40" S
145° 25' 30" E), badly stock-damaged, on
Edgbaston Station in 1988. A second lot of nine
specimens came from the “Western” spring,
about 2.4 km NE of the station homestead. 'Hie
pools and springs are fed by Artesian Basin
water (Ivantsoff et al. 1991). Ivantsoff et al.
(1991), in describing the habitat of a new
pseudomugilid, indicated that Chlamydogobius
shared its habitat, which consists of small, clear,
very shallow, clay-bottomed pools, with grass
tussocks growing in the water and an unidenti¬
fied red-leafed plant.

There has been concern expressed (Unmack
1992) that the pools on Edgbaston Station may
become degraded by trampling (sheep, cattle
and human beings), and that the fish populations
may become more directly threatened by the
introduced pest fish, Gambusia holbrooki , which
is already present in a number of the pools,
including those with Chlamydogobius present.
The status of the population of C. squamigenus
isnot known, although Unmack (1992) indicates
that there were populations in several pools. The
ASFB Threatened Fish Committee lists this spe¬
cies as “vulnerable” (Jackson 1993).

Bruce Grose (in litt.) has been able to spawn
and raise this species in captivity, and reports
that the eggs take about a week to hatch into free-

H.K. Larson

swimming larvae which actively disperse through
the aquarium. He estimates at least a hundred
eggs per batch are laid.

Remarks. Ivantsoff et al. (1991) refer to this
species as “... possibly a new species of the desert
goby” in their description of the unusual
pseudomugilid Scaturiginichthys, but do not dis¬
cuss the goby further.

Crossmoor Bore specimens included two with
vertebral counts of 11+17 and two with dorsal
pterygiophore patterns of 3-11210.

Etymology. From the Latin squama (scale)
and gena (cheek), referring to the cycloid scales
present on the preopercular region of this spe¬
cies. All other species of the genus lack scales on
the preopercle.

PALAEOBIOGEOGRAPHY

The present Australian freshwater fish fauna
is well known for its several ancient Gondwanan
relicts such as Neoceratodus and Scleropages,
which are now restricted to tropical/sub-tropical
areas. However, the origins of the remaining
marine-derivedfauna is not known (Allen 1989).
The present temperate (southern) freshwater fish
fauna does include a number of families of
probable Gondwanan coastal marine ancestry
(Aplochitonidae, Galaxiidae, Prototroctidae,
Retropinnidae, Percichthyidae, Gadopsidae);
most of these families include species occurring
in southern Australia, New Zealand and South
America. The present northern and central fresh¬
water fish fauna includes groups with SE Asian
affinities (Plotosidae, Melanotaeniidae,
Ambassidae, Terapontidae, Gobiidae). Allen
(1982) speculated that many Australian fresh¬
water fish species may possibly have evolved
only in the last 2 my. Allen and Cross (1982)
state that rainbowfish (family Melanotaeniidae,
endemic to Australia and New Guinea) are capa¬
ble of speciating rapidly (7,000 years estimated
for the M. trifasciatalM. goldei species pair; the
former is from northern Australian, the latter
from southern New Guinea).

Fossil gobiids have been known since the
Eocene, based on otoliths (Miller 1973), and are
known from the famous Monte Bolca site (mid¬
dle Eocene) which includes the first records of
many modem coral reef groups such as labrids,
pomacentrids, zanclids, acanthurids siganids and
ephippids (Bellwood 1993). Miller considered
that the other gobioids may have split from the
rhyacichthyids in the late Cretaceous, when many

teleost fish families appeared. There are no fossil
gobioids known so far from Australia (Long and
Turner 1984).

Larson (in prep.) and Miller (1987) both con-
siderthattheclosestrelativeofC/i/amyrfogoi/nA'
may b eMugilogobius, an estuarine genus related
to other Indo-West Pacific genera such as
Pseudogobius and Hemigobius. These fish are
likely to have radiated from the SE Asian Archi¬
pelago viaNew Guineaalongwithothershallow-
water fish groups to the Australian coast. Miller
(1987) has postulated that Chlamydogobius be¬
came isolated in central Australia approximately
1.6 my, after enteri ng the continent about 20 my.
To determine how Chlamydogobius entered cen¬
tral Australia and radiated, one must depend upon
hypotheses of phylogenetic relationships,
palaeogeographic recontructions, physiological
tolerances of the fish, and climate indicators,
rather than a fossil record. In Australia, there is
only one poorly known Palaeogene (65-22.5 my)
vertebrate fossil deposit, while the Neogene
(Miocene and Pliocene, 22.5-1.8 my) and
Pleistocene (1.8 my to 10,000 years) faunas are
betterknown (Rich 1991). Long (1991) indicated
that many modem genera of shallow-water ma¬
rine fishes such as Lactarius, Diodon, Sillago and
Platycephalus were established in Australia dur¬
ing theearly Miocene, but this should be taken as
a minimum age because there is no older fish
record with which totime their arrival (Megirian
pers. comm.).

Based on available information (e.g. Frakes
and Rich 1991; Megirian 1992; McGowran and
Li 1994), the early to mid Miocene were possibly
optimum times forthe introduction and dispersal
of gobiids into the interior (in general agreement
with Miller 1987). During the “Mioceneoscilla¬
tion” (McGowran and Li 1994), sea surface
temperatures and sea levels rose and some tropi¬
cal marine groups were able to spread around the
southern coast of Australia. Inland, conditions
also ameliorated, but were still relatively dry
(Megirian 1992). However, there was probably
significantly more permanent water inland than
there is today, fresh grading to brackish in en¬
closed or endorheic basins. The general low
relief of most of the continent was established at
least since the Early Cretaceous (e.g. Fig. 18 in
Frakes and Rich 1991 shows much of the conti¬
nent flooded, indicating its flatness); many ma¬
jor drainages were probably already separated
by divides of very low relief which probably
would not have formed barriers to dispersal
during times of flood. The low drainage divide

A review of the Australian endemic gobiid genus Chlamydogobius

between the (present-day) Gulf and Lake Eyre
drainages may have been present since the early
Cretaceous (Swarko 1966), or Palaeocene
(Veevers 1991).

There are two possiblepaths thatthe ancestor(s)
of present-day Chlamydogobius could have taken
to enter central Australia: along coastal New
Guinea into the Gulf drainage and thus into the
Lake Eyre basin (where radiation intocentralian
drainages could occur), or along coastal New
Guinea, around the eastern coast of Australia to
the southern margin, then north into the Lake
Eyre basin andcentralian drainages. The former
scenario requires a connection between the Gulf
and Lake Eyre drainages. The old but probably
low drainage division between the two may have
been breached during one of the wetter climatic
periods. Given the present-day ability of many
gobioids (such as the world-wide subfamily
Sicydiinae, the Indo-West Pacific genus
Glossogobius, the Australia-New Guinea genus
Mogurnda, and Gobiomorphus of New Zealand
and south-eastern Australia) to colonise high-
gradientfaststreamsand/orclimbnearly vertical
surfaces, and the present distribution of C. ra¬
nunculus, this scenario is not unlikely.

Whichever route was used, Chlamydogobius
must have been established in central Australia
before the arid conditions of today were estab¬
lished, as could have other genera such as
Neosilurus and Craterocephalus , which have
also speciated in the interior (it is interesting to
note that central Australian fishes are a mixture
of widely distributed, opportunistic species, such
as Leiopotherapon unicolor and Nematalosa
erebi, or those with relatively restricted distribu¬
tions such as the species of Chlamydogobius,
Craterocephalus centralis and two undescribed
Neosilurus). The aridification of the interior to
the extent observed today began in the late
Miocene/early Pliocene, with the formation and
mobilisation of desert dune systems first occur¬
ring in the late Pliocene and becoming wide¬
spread in the Pleistocene, especially during gla¬
cial epochs, as a result of global temperature
gradients, formation or intensification of sub¬
tropical high-pressure belts and consequent in¬
creases in climatic seasonality in the continental
interior. The Finke drainage system may have
been isolated quite recently from the Lake Eyre
basin by dune development and migration in the
Pleistocene (Megirianpers. comm.), which would
seem consistent with the small genetic differ¬
ence exhibited by Chlamydogobius eremius and
C. japalpa (as observed by Adams). Thus, the

intensifying aridification of Australia led to iso¬
lation of Chlamydogobius populations and sub¬
sequent speciation.

DISCUSSION

Preliminary work by Mark Adams at the South
Australian Museum indicated that there were
possibly four distinct freshwater species of this
genus, based on isozyme electrophoresis (Terry
Sim and Mark Adams, pers. comm.). Mark Adams
conducted enzyme electrophoresis on five inland
populations of Chlamydogobius and considered
that four of them were probably valid (Adams, in
prep.). He did not consider that there was suffi¬
cient difference between the Finke and Coward
Springs populations to warrant placing them in
separate species (consensus analysis of his data
indicated no genetic difference, although one
locus, Ck, showed as a separating factor depend¬
ing on how the data was input into the clustering
procedure). He found considerable difference
between C. gloveri and C. eremius, reflecting the
isolation between the Lake Eyre system and
Dalhousie Springs as proposed by Kotwicki
(1989). The 1 i nk between the Lake Eyre and Finke
systems is supported by Crowley and Ivantsoff
(1990b), who found an allele shared by
Craterocephalus centralis (restricted to the Finke
system) and C. eyresii (Lake Eyre drainage) wh ich
was absent from others in their species-group.
Given the long-term hydrologic isolation of the
Finke system from Lake Eyre, and the morpho¬
logical and axial skeleton differences between the
two populations of Chlamydogobius, they are
treated here as separate species. As Ivantsoff,
Larson and Ponder each note (in Department of
the Environment, Sport and Territories 1994:49),
more species of freshwater fish are being found
to be narrowly restricted in theirdistributions, i.e.
found only in a particular group of springs.

As Miller (1987) noted, Chlamydogobius ex¬
hibits several adaptive features which contribute
to its survival in hostile environments: a wide
tolerance range for temperature, pH, salinity and
oxygen levels, and large egg size with almost no
larval stage, the young fish clinging almost im¬
mediately to the substrate; the latter two traits
differing from Mugilogobius (Horsthemke 1989;
Armstrong 1994 pers. comm.), which has a short
pelagic larval stage. Glover (1982) discussed the
physical barriers operating which influence desert
fish dispersal and present-day distribution, and
showed that the ability to tolerate extremes of

H.K. Larson

Plate 1 , above, Chlamydogobius eremius, male in breeding condition. Photo by Neil Armstrong;
center, Chlamydogobius eremius, male in non-breeding coloration. Photo by Ross Felix;
below, Chlamydogobius gloveri n. sp., female. Photo by Ross Felix.

A review of the Australian endemic gobiid genus Chlamydogobius

Piute 2, above, Chlamydogobius gloveri n. sp.. male. Photo by PeterUnmack; center, Chlamydogobius
ranunculus n. sp., adult male. Photo by Rex Williams; below, Chlamydogobius ranunculus n. sp.,

adult female. Photo by Neil Armstrong.

H.K. Larson

temperature and salinity, as well as flexibility in
handling other ecological factors, favour sur¬
vival for fishes such as the desert goby,
Chlamydogobius.

ACKNOWLEDGMENTS

Many thanks to those who kindly provided
specimens, colour slides and/or information, es¬
pecially die late John Glover, also Mark Adams,
Neil Armstrong, Ross Felix, Bruce Grose, Paul
Homer, Richard Kimber, Keith Martin, Terry
Sim, Peter Unmack, Rob Wager and Dave
Wilson. Terry Sim made available John Glover’s
original data on a number of Chlamydogobius
populations. I am indebted to Dirk Megirian
(NTM) for detailed discussion and information
on palaeogeography and geologic history. Rex
Williams drew the maps, and he and Rick Bawden
photographed the type specimens.

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Accepted 20 June, 1995

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:53-59

LARVA, PUPA AND NOTES ON GENERAL BIOLOGY OF TINODES
RADONA NEBOISS (TRICHOPTERA: PSYCHOMYIIDAE).

A. WELLS

Australian Biological Resources Study,
GPO Box 636, Canberra, ACT 2601, Australia.

ABSTRACT

The larva and pupa of the north Australian psychomyiid caddisfly Tinodes radona
Neboiss, 1990, are described and figured for the first time. In addition, the larval habitat
is illustrated and the pupal habitat is noted and discussed, particularly in relation to the
unusual pupal mandibles.

Keywords: Trichoptera, Psychomyiidae, Tinodes radona, larva, pupa, habitat.

INTRODUCTION

The family Psychomyiidae was reinstated re¬
cently on the list of Australian Trichoptera with
the description by Neboiss (1990) of two species
from northern Australia. Previously, by elevat¬
ing the subfamily Ecnominae to family status,
Neboiss (1977) had removed the record of the
family from the Australian fauna. Since the
Psychomyiidae sensu stricto (excluding the
Xiphocentronidae) is widespread in the world,
being noted by Wiggins (in 1982, and therefore
prior to Neboiss’ rediscovery for Australia) as
absent only from the Neotropics and Australia, it
was not surprising to have representatives of the
family found in Australia.

One of the two Australian species, Tinodes
radona Neboiss, 1990, is based on descriptions
of male and female; for the other, Zelandoptila
yuccabina Neboiss, 1990, only the male is de¬
scribed. Larvae and pupae of Tinodes radona are
now known. They are similar in aspects of mor¬
phology and habits to immatures of some
Holarctic species in several genera of the
Psychomyiidae and its sister-group, the
Xiphocentronidae. However, for completeness
of records of the Australian Trichoptera fauna,
larval and pupal features and aspects of their
general biology are described and illustrated
here. As yet, nothing is known of immatures of
Z. yuccabina.

Neboiss’ (1990) work on T. radona was
based on specimens from the northern part
of the Northern Territory, colloquially the
‘Top End”, and from Cape York Peninsula,
north Queensland. Additional adults have
now been collected from both regions, and in
addition, larvae and pupae have been col¬
lected from several ‘Top End” localities.

Adult Tinodes radona generally have been
taken at lights or in light traps, but several recent
Northern Territory collections were made by
sweep-netting of vegetation overhanging and
bordering waterfalls and cascades. With consid¬
erable delight, when collecting in this way on
one occasion, I realised that the “chironomid”
tubes commonly seen in the splash zone of falls
and cascades (Plate 1 a) and above water level on
boulders (Plate lb) in streams in the “Top End”,
are, in fact, the silken retreats of T. radona
larvae. Larvae of Tinodes are reported to feed
mainly on detritus and algae (Wiggins 1977)and
presumably extend their silken tubes as they
graze on the algal material on the rock surface.
The finding of larvae, and subsequent searches
for pupae, led eventually to the discovery of the
sand grain pupal cases on damp rock above water
level close to the larval tubes.

The T. radona larva and pupa described and
figured below were taken from the micro-habitat
illustrated in Plate lb, in Kakadu National Park,
Northern Territory. All specimens are in the

A. Wells

Figs 1-6. Tinodes radona Neboiss larva: 1,2 head, ventral and dorsal views; 3, mandibles, dorsal view; 4, labrum,
dorsal view; 5, anal claw; 6, habitus, lateral view. Abbreviations: a.v.a., anterior ventral apotome; sm. s.,
submental sclerite. Scale bars 0.1 mm, unless otherwise indicated.

North Australian caddisfly

collection of the Northern Territory Museum of
Arts and Sciences, Darwin, together with other
specimens collected from the hygropetric zone
of a small waterfall in Litchfield National Park,
Northern Territory. Slide mounts were prepared
for study by macerating specimens in caustic
potash, clearing them in clove oil and subse¬
quently mounting them in Canada balsam.

DESCRIPTION

Material examined. 2 larvae, NT, Kakadu
National Park, Baroalba Creek, above Koobarra
Pools, 4 October 1991, A. Wells; numerous
larvae, same locality and collector, 28 June 1992;
numerous larvae and pupae, same locality and
collector, 12 September 1992; 2 larvae, NT,
Litchfield National Park, Aida Creek, 24 June
1992, A. Wells; 1 pupa, Litchfield National Park,
Tjaetabe Falls, 4 September 1992, A. Wells and
J. Webber.

Larva, final instar (Figs 1-8). Body elongate,
vermiform; length about 4.6-6.7 mm. Head and
thoracic sclerites dark brown. Head in dorsal
view (Fig. 2), slightly longerthan wide, rounded
laterally, truncate anteriorly and posteriorly.
Frontoclypeus with anterior margin almost trun¬
cate, without setae. Ventrally on head (Fig. 1), a
row of setae near anterior and posterior lateral
angles, posterior setae very short, one seta beside
median suture at about 2/3 length; anterior ven¬
tral apotome short, crescentic; posterior ventral
apotome absent; paired sub-mental sclerites
subquadrate. Labrum (Fig. 4) short, rounded
anteriorly, with dense short setae antero-later-
ally. Mandibles (Fig. 3) short, triangular in dor¬
sal and ventral views, without clear denticles and
with pair of elongate setae laterally; left mandi¬
ble with well-developed penicillus; right mandi¬
ble with small tuft of setae towards base on inner
dorsal margin. Labium forming long slender
spinneret (Fig. 1).

Thorax with only pronotum sclerotised, nar¬
row in comparison to meso- and meta-nota.
Foretrochantin broad, separated from pro-pleuron
by clear suture (Fig. 8). Legs short; forecoxa
subquadrate (Fig. 7); tarsal claws rounded. Ab¬
domen without gills. Anal pro-legs short, robust,
darkly sclerotised, strongly hooked (Fig. 5). Anal
papillae absent.

Pupa (Figs 9-15). Length about 3.0-3.5 mm.
Labrum (Fig. 12) with anterior margin entire,

broadly rounded, with clustered setae apico-lat-
erally and at baso-lateral margin, short setae
mesially. Mandibles (Fig. 13) elongate, length
almost 3X basal width, tapered to fine straight
section subapically, a small cluster of
denticles apically, directed posteriorly; paired
lateral setae towards base. Anterior and pos¬
terior hookplates on abdominal segment V,
anterior hookplates only on segments II-IV
and VI-VIII (Fig. 14). Abdomen with short,
tapered apical processes (Fig. 15), setae clus¬
tered at apices.

Pupal case (Figs 9, 10), length 4.2-6.5 mm,
conical in shape, built of coarse sand grains, flat
ventrally over area of contact with substrate, a
perforated, more or less ovoid closure mem¬
brane (Fig. 11) on ventral surface at broadest
(anterior) end.

Habitat (Plate la-b). Silken larval tubes up to
30 mm long are found on damp rocks above
water at the edges of or in small streams (Plate
lb), and in the splash zone beside waterfalls,
cascades (Plate 1 a) and riffles, in shaded or open
sunny areas. Pupal shelters are constructed
on damp rock, above or adjacent to flowing
water; the shelters are generally separated
by a short distance from the larval tubes.
Adults are diurnal and can be seen by day
beside flowing water, on rocks and riparian
vegetation.

Figs 7, 8. Tinodes radona Neboiss larva: 7, forelimb;
8, prothoracic pleurites.

A. Wells

Plate 1. Larval tubes of Tinodes radona Neboiss: a. (top), on rocks in splash zone of a cascade, Oenpelli, Arnhem Land,
NT (coin diameter =28 mm); b, (below), on a rock in midstream above Koobarra Springs, Kakadu National Park, NT.

North Australian caddisfly

Figs 9-11. Tinodes radona Neboiss pupal case: 9-10, dorsal, ventral views', 11, closure membrane.
Figs 12-15. Tinodes radona Neboiss pupa: 12, labrum; 13, mandibles; 14, hookplates; 15, terminal
abdomen. Scale bars 0.1 mm, unless otherwise indicated.

A. Wells

Remarks. Larvae of Tinodes radona are eas¬
ily distinguished from other Australian species
by the combination of membranous meso- and
meta-nota and the broad, blade-like fore-
trochantin. Pupae are recognised by the shape of
their mandibles and hookplates.

DISCUSSION

A survey of the literature on the Psychomyiidae
re veals th at at least one B ri tish species of Tinodes,
T. assimilis McLachlan, is foundin microhabitats
similartothatofr. radona (Hickin 1967).IIickin
(1967) also mentions that larvae of several other
Tinodes species are able to withstand removal
from free water, eitherby a drop in water level or
when collected for study, for considerable peri¬
ods of time. In North America (Wiggins 1977),
and Europe (Lepneva 1964)the larvae of Tinodes
appear to live submerged in water, but some
larvae in one of the psychomyiid sister group, the
Xiphocentronidae, are reported to be hygropetric
(i.e. Xiphocentron-see Edwards 1961; Flint 1964;
Wiggins 1977).

A curious feature of both the Psychomyiidae
and Xiphocentronidae is their modified pupal
mandibles. These are illustrated for the Euro¬
pean Tinodes rostocki McLachlan by Lepneva
(1964, but her figures are taken from another
author), for Xiphocentron ijaitiensis (Banks) by
Flint (1964), and here for T. radona (Fig. 13).
Edwards (1961) shows a more extreme ver¬
sion of these mandibles in X. mexico Ross, in
which the distal parts of the mandibles are
whip-like and Hickin (1967) mentions “long
scythe-like pupal mandibles” in the
psychomyiid Lype reducta (Hagen). Mem¬
bers of the Polycentropodidae and
Ecnominae have elongate pupal mandibles
but without the distal hook. Since other caddis
species that build sand pupal shelters have
pupae with unmodified mandibles, it is pos¬
sible that the curiously hooked mandible
evolved with the adoption of an unusual
niche. Examination of the emergence be¬
haviour of these pupae could be rewarding.

In normal situations, immersed in water,
pharate adult caddisflies escaping from their
pupal enclosures would be well buoyed up by the
surrounding water; pupae removed from their
cases into free water often prove hard to “sink",
tending to float on the surface of the collecting

vessel. However, pharate adults of T. radona,
when emerging in the hygropetric zone, presum¬
ably drag themselves, unaided by buoyancy of
water, from their sand grain case, against
resistance from the substrate and particu¬
larly from its “aufwuchs” covering. The pupa]
mandibles, which show close resemblance
in form to the spagetti scoop of a modern
kitchen, probably aid the exit from the case
and may also allow the pharate adult to gain
some hold in the fine mat on the substrate
prior to emergence from the pupal cuticle.

ACKNOWLEDGEMENTS

Specimens were collected in Kakadu National
Park with permission from the Australian Na¬
tional Parks and Wildlife Service (now Austral¬
ian Nature Conservation Agency), and from
Litchfield National Park with permission of die
Conservation Commission of the Northern Ter¬
ritory. The work was funded by the Northern
Territory Museum of Arts and Sciences. Jenni
Webber assisted with field work and final prepa¬
ration of the plates; Ian Morris and Phillip
Kempster photographed the larval tubes. Com¬
ments by David Cartwright and two referees
greatly improved the manuscript. I wish to thank
the above mentioned and also staff of the Office
of the Supervising Scientist at Jabiru for gener¬
ous assistance and encouragement with work in
the Alligator Rivers region. The manuscript was
prepared using facilities provided by the Aus¬
tralian Biological Resources Study and the
Division of Entomology, CSIRO, Canberra.

REFERENCES

Edwards, S.W. 1961. The immature stages of
Xiphocentron mexico (Trichoptera). Texas Jour¬
nal of Science 13: 51-56.

Flint, O.S. 1964. The caddisflies (Trichoptera) of
Puerto Rico. University of Puerto Rico, Agricul¬
tural Experiment Station, Technical Paper 40: 1-
80.

Hickin, N.E. 1967. Caddis larvae. Hutchinson: Lon¬
don.

Lepneva, S.G. 1964. Fauna SSSR, Rucheiniki, vol. 2,
no. 1. Lichiniki i kukolki podotryada
kol’chatoshchupikovykh. Zoologicheskii Institut
Akademii Nauk SSSR, n.s. 95. [In Russian.
Translated into English as: Fauna of the U.S.S.R .;

North Australian caddisfly

Trichoptera, vol. 2, no. 1. Larvae and Pupae of
Annulipalpia. Published by the Israel Program
for Scientific Translations, 1971].

Neboiss, A. 1977. A taxonomic and zoogeographic
study of Tasmanian caddis-flies (Insecta:
Trichoptera). Memoirs of the National Museum
of Victoria 38: 1-208.

Neboiss, A. 1990. The family Psychomyiidae
(Trichoptera) re-established in Australia. Mem¬
oirs of the Museum of Victoria 51: 83-86.

Wiggins, G.B. 1977. Larvae of the North American
caddisfly genera (Trichoptera). University of
Toronto Press: Toronto and Buffalo.

Wiggins, G.B. 1982. Trichoptera. In: Parker, S. (ed.)
Synopsis and Classification of Living Organ¬
isms. Pp. 599-612. McGraw-Hill: New York.

Accepted 15 February, 1994

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:61-64

LATREUTES AN OP LON YX KEMP, 1914 (CRUSTACEA: DECAPODA:
HIPPOLYTIDAE), A JELLY-FISH ASSOCIATE NEW TO
THE AUSTRALIAN FAUNA.

A.J. BRUCE

Division of Natural Sciences, Museum and Art Gallery of the Northern Territory,
P.O. Box 4646, Darwin, NT 0801, Australia.

ABSTRACT

This report records the presence of the hippolytid shrimp Latreutes anoplonyx Kemp
in the Australian fauna, from the Northern Territory and Queensland, and confirms
beyond doubt that the species is a genuine associate of scyphozoans.

Keywords: Latreutes anoplonyx, Crustacea, Caridea, Hippolytidae, new to Australian
fauna, association with Scyphozoa confirmed.

INTRODUCTION

Few caridean shrimps have been reported as
associates of scyphozoancoelenterates. One of the
first observations of possible association was of
the hippolytid shrimp Latreutes anoplonyx Kemp,
noted by De Man (1929), but the basis of the
observation was the discovery of a shrimp “in” a
medusa, and not on the study of live shrimps and
their host in vivo. The present observations appear
to be the first of a live hippolytid shrimp in asso¬
ciation with jelly-fish hosts.

DESCRIPTION

Latreutes anoplonyx Kemp, 1914

Restricted synonymy

Latreutes anoplonyx Kemp, 1914: 104-105,
pi. 4, figs 3-5;- 1916: 369.

Material. 3 females (2 ovig.). Van Diemen
Gulf, NT, 36 m, 6 June 1982, FV Skelton, NTM
Cr.0000191.22 spms (6 ovig. females), Cameron’s
Beach, Darwin, NT, 9 March 1978, coll.
N.T.Fisheries, NTM Cr.0000191. 28 spms (21
ovig. females), Auckland Creek, Gladstone, Qld.,
8 January 1982, coll. P. Saenger, NTM
Cr.0002640. 16 spms (no ovig. females), Cham¬
bers Bay, Van Diemen Gulf, NT, May 1978, coll.
J. Grice, NTM Cr.0007418. 1 male, 1 female,
from inner part of Darwin Harbour, about 4.8 km
from the wharfs, NT, coll. T. Heeger, 4 June 1993,
NTM Cr.010640. 2 males, 2 females, 1 juv..

mouth of Buffalo Creek, Darwin Harbour, NT, 12
March 1993, 0.75 m, coll. P.N. Alderslade, P.

Homer and N. Smit, NTM Cr.0010531.

Comparative description. The specimens
agree well with the descriptions providedby Kemp
(1914, 1916) and Liu (1955), but appear to differ
slightly from that figured by Hayashi and Miyake
(1968b). The adult Darwin Harbour specimens
have 12-14 small, acute, dorsal rostral teeth, with
9-10 similar ventral teeth (Fig. 1). The dentate
dorsal margin is feebly concave and the ventral
margin straight, with the rostral process appearing
acute in lateral view. The proximal dorsal rostral
tooth is of similar size to the rest of the series, in
contrast to the material from the Amakusa Islands,
in which it appears distinctly larger than the rest of
the series. The rostrum also appears much deeper
and less acute, with the distal dorsal border straight
and not concave. In the specimens illustrated by
Liu, the rostrum also appears much deeper than in
the Australian specimens, and in his ovigerous
female both dorsal and ventral margins appear
distinctly convex. The dactyls of the ambulatory
pereiopods are less strongly spinose than those
illustrated by De Man (1929). The ventral margin
of the corpus bears only two small spinules, in¬
stead of four as shown by De Man and described by
Hayashi and Miyake (1968a), but also noted with
two spinules (Hayashi and Miyake 1968b) for the
fourth pereiopod. The unguis is comified and only
feebly distinct from the corpus.

Measurements. The largest adult female speci¬
men from Darwin Harbour has a postorbital cara¬
pace length of 13.5 mm and a total body length of

A. J. Bruce

27.0 mm. The largest specimens were from
Gladstone, with a postorbital carapace length of
21.9 mm and a total body length of 47.0 mm.

Colouration. The colour pattern of the live
shrimps (Fig. 2), not previously reported, closely
matches that of the host scyphozoan and is clearly
of a cryptic nature. The general colouration is a
dull reddish brown. The distal half of the rostrum
is a deeper red, with yellow dots proximally; the
proximal half colourless. The carapace has a nar¬
row transverse white bar on the anterior cardiac
region, curving anteroventrally to fade out on the
central branchiostegite. Anteriorly, the carapace
is dark red dorsally, with large white spots;
posteriorly the carapace is pale red, with scattered
darker spots; extending, on to the lower
branchiostegite. The abdominal segments are simi¬
larly coloured, each having a narrow transverse
white bar extending onto the central or lower
pleuron, with a narrow, dark red band posteriorly
and a broader, paler area anteriorly; the posterior
two-thirds of the sixth segment and the caudal fan
are colourless. The proximal part of the rostrum,
antennule and antenna, and the second to fifth
pereiopods are similarly colourless although the
meri of the ambulatory pereiopods are feebly spot¬
ted with red. The basipodites of the pleopods are
similarly speckled with red. The basicerite, third

maxilliped and first pereiopod are all reddish. The
cornea is pallid. The colour pattern is less cop,
spicuous in the smaller males.

Hosts. Phyllorhiza punctata von Lendenfefr,
(Mastigiidae), and Versuriga anadynomene
(Maas), (Versurigidae) [Coelentcrata: Scyphozoa].
The associations with Phyllorhiza and Versurig#
represent new host records.

Behavior. The Buffalo Creek shrimps were
collected from two specimens of the jelly-fish ho$t
obtained by bucket from surface water over 0.75 pi
depth. The larger jelly-fish had a bell diameter of
about 150 mm, and the shrimps were first seep
swimming between hosts in the container. The
inner Darwin Harbour specimens were collected
from a surface swimming specimen of Versuriga,
over water of uncertain depth. They were observed
alive, moving actively around over the
subumbrellar surface and rather lethargically over
the tentacles.

Distribution. The species was first described
by Kemp (1914) from specimens from Bombay,
with subsequent specimens from Bombay and
from off the coast of Myanmar, at a depth of 13-
15m. Further records have been from Indonesia
(De Man 1929), Tanjung Krawang, east of Ja¬
karta, Java; (Holthuis 1947), Jakarta and off
Panaroekan, eastern Java; China (Liu 1955),

Fig. 1 . Latreutes anoplonyx Kemp, Buffalo Creek. Darwin. Anterior carapace and rostrum. A, male; B, smaller female, tip
of rostrum; C, larger female; D, male; E, dactyl of third pereiopod, female.

Shrimp associate of jelly-fish

Fig. 2. Latreutes anoplonyx Kemp. A, female, Buffalo Creek, Darwin; B, male and female, Darwin Harbour; C, female,
Darwin Harbour. Photographs by A. J. Bruce.

A. J. Bruce

Liaoning, Hopei, Shantung and Kiangsu prov¬
inces; Singapore (Johnson 1961); Japan (Hayashi
and Miyake 1968a), Tanabe Bay; ( Hayashi and
Miyake 1968b), Amakusa Islands.

Remarks. The possible association of this spe¬
cies with scyphozoan hosts was first suggested by
De Man (1929), whose specimens, three females,
were found “in” a jelly-fish, a specimen of
Rhizostoma (?). One of Holthuis’ (1947) speci¬
mens was found in the subgenital cavity of
Acromitus flagellants. The specimens of Liu
(1955) were associated with Rhopilema esculenta.
Hayashi and Miyake (1968a) reported an associa¬
tion with Mastigias papua. Details of the collec¬
tion of these specimens were not provided and it is
likely that the material was obtained from trawl
catches, with the possibility that the associations
were accidental, the result of the shrimps and the
jelly-fish sharing the same habitat. There appear to
have been no observations on the shrimps in life.
The specimens here described were collected by
hand, with a bucket, in the course of a survey for
shallow water toxic cubomedusae, and were im¬
mediately observed on the host. The col our pattern
of the shrimps closely resembles that of the host
animals, which are also heavily mottled with brown
patches.

DISCUSSION

Associations between shrimps and jelly-fish are
uncommon. Hayashi and Miyake (1968a) sug¬
gested that Chlorotocella gracilis, a pandalid
shrimp, and Latreutes mucronatus were also jelly¬
fish associates. The real habitat of the former
species seems rather uncertain although it is quite
common in tropical waters. It is frequently found
in numbers in association with Periclimenes
psamathe De Man on gorgonian hosts, as well as
in plankton samples taken at night Latreutes
mucronatus is a common shrimp in sea grass beds
and is particul arly like ly to be col lected when jel ly-
fish are sampled by trawl. Two pontoni ine shri mps
have been reported as jelly-fish associates, both on
the basis of direct underwater observation.
Periclimenes holthuisi is frequently associated
with the sedentary shallow water Cassiopea spe¬
cies (Bruce 1972). The only known specimen of
Periclimenes tonga was also observed in situ on
its unidentified host jelly-fish (Bruce 1989).

The specimens from FV Skelton were caught by
trawl from water with a depth of 36.6 m, the
greatest depth from which this species has been
recorded. It is possible that, as few examples were
obtained, these specimens may have been from a
jelly-fish caught near to the water surface. The

other specimens were probably from shallow wa¬
ter and their largernumbers suggest that they were
collected from the bottom.

ACKNOWLEDGEMENTS

I am most grateful to Mr P.N. Alderslade and Dr
T. Heeger for the specimens of Latreutes and the
identification of their scyphozoan hosts.

REFERENCES

Bruce, A.J. 1972. An association between a
pontoniinid shrimp and a rhizostomatous
scyphozoan. Crustaceana 23(3): 300-302.

Bruce, A.J. 1989. Periclimenes tonga sp. nov., a
commensal shrimp associated with a scyphozoan
host from Tonga (Crustacea: Decapoda:
Palaemonidae). Micronesica 21(1988): 23-32.
Hayashi, K.-I., and Miyake, S. 1968a. Three caridean
shrimps associated with a medusa from Tanabe
Bay, Japan. Publications from the Seto Marine
Biological Laboratory 16: 11-19.

Hayashi, K.-I., and Miyake, S. 1968b. Studies on the
hippolytid shrimp fauna of Japan, V. Hippolytid
fauna of the sea around the Amakusa Marine
Biological Laboratory. Ohmu, Occasional Pa¬
pers of the Zoological Laboratory, Faculty of
Agriculture, Kyushu University, Fukuoka, Ja¬
pan 1(6): 1212-163.

Holthuis, L.B. 1947. The Decapoda of the Siboga
Expedition. X. The Hippolytidae and
Rhynchocineudae collected by the Siboga and
Snellius Expeditions with remarks on other spe¬
cies. Siboga Expedition Monograph 39a(8): 1-
100 .

Johnson, D.S. 1961. Prawns of the Malacca Straits
and Singapore waters. Journal of the Marine
Biological Association of India 18(1): 1-54.
Kemp, S. 1914. Notes on Crustacea Decapoda in the
Indian Museum. V. Hippolytidae. Records of
the Indian Museum 10: 81-129.

Kemp, S. 1916. Notes on the Crustacea Decapoda in
the Indian Museum. VII. Further notes on
Hippolytidae. Records of the Indian Museum
12: 385-405.

Liu, J.Y. 1955. Economic shrimps and prawns of
northern China. Marine Biological Institute,
Academy of Sciences, Peking, pp.i-iii, 1-73, pis
1-24 (in Chinese).

Man, J.G.de. 1929. On a small collection of Decapoda,
one of which a Crangon, caught by the Danish
Pacific Expedition at the Jolo Islands, is new to
science. Papers from Dr.Th. Mortcnsen’s Pa¬
cific Expedition 1914-1916. Videnskabelige
Meddelelser fra Dansk naturhistorisk Forening
i Kobenhavn 87: 105-134.

Accepted: 31 January, 1994

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:65-74

NOTES ON THE GENUS ANISOPS SPINOLA (HEMIPTERA-
HETEROPTERA, NOTONECTIDAE) OF THE NORTHERN TERRITORY

AND WESTERN AUSTRALIA.

I. LANS BURY

Hope Entomological Collections, University Museum,
Oxford 0X1 3PW, England.

ABSTRACT

A sample of water-bugs from the Elkedra River, about 340 km north east of Alice
Springs, Northern Territory, has a remarkable mixture of Notonectidae, Anisops
Spinola species from two zoo-subregions (Torresian and Eyrean), including a new
species Anisops elkedraensis. Additionally, Anisops hayesi sp. nov. is described from
the Alice Springs area and Anisops ungarinyin sp. nov. from the central Kimberleys,
Western Australia.

Keywords: Hemiptera, Notonectidae, Anisops, new species, Western Australia,
Northern Territory.

INTRODUCTION

Amongst the collections of water-bugs in the
Northern Territory Museum, Darwin, a sample
from the Elkedra River (about 340 km north¬
east of Alice Springs), 21°13S,135° 085’E, 28
September 1983, collected by P. Horner, H.
Larson and K. Bishop, has been found to have an
unusually high diversity of Anisops spinola spe¬
cies which includes Torresian and Eyrean ele¬
ments. Two species, Anisops gratus Hale and A.
thienemanni Lundblad, also extend over much
of the Bassian region and are compared with
forms from southern Queensland and south¬
western Australia respectively. The forms are
figured and the possibility that A. thienemanni
may be a group of sibling species is discussed
briefly. Anisopsparacrinita Brooks and A.nasuta
Fieberare listed. Included in these notes are the
descriptions of Anisops hayesi sp. nov. from
Alice Springs and Anisops ungarinyia sp. nov.
from north-western Australia.

The Elkedra specimens including the type
series of Anisops elkedraensis sp. nov. are de¬
posited in the Northern Territory Museum
(NTM), Darwin.

SYSTEMATICS

Family Notonectidae Leach
Anisops gratus Hale
(Figs 1-7)

Anisops gratus Hale, 1923: 413-414;-Brooks
1951: 352-353; -Sweeney 1965: 91; -Lansbury
1969: 448-449.

Material examined. NTM Entomological col¬
lection, Northern Territory, Elkedra River, 1 male.

Description. Male 8.54 mm long, 2.32 mm
wide. Elkedra male: chaetotaxy of front leg (Fig.
1), stridulatory comb (Fig. 2), rostral prong (Fig.
4). Head and pronotum (Fig. 3), note long
pronotum. This male has been compared with a
series from south Queensland, Wyara System,
Werewilka Creek (Wyara System on Boorara
Station, Qld), total dissolved solids 12.8 g/1,
coll. B.V. Timms. Male front leg (Fig. 5),
stridulatory comb (Fig. 6), rostral prong (Fig. 7).
The rostral prong of the Queensland form is
rather more sharply curved and the facial tuber¬
cle is slightly raised.

Distribution. The distribution of A. gratus is
predominantly Eyrean and Bassian; the record

I. Lansbury

Figs 1-4. Anisops gratus male (Elkedra): 1, front leg, inner view, slide mounted; 2, stridulatory comb, slide mounted; 3, head
and pronotum, dorsal view; 4, rostral prong. Scale line 0.5mm.

Figs 5-7. Anisops gratus male (Wyara): 5, front leg, inner view, slide mounted; 6, rostral prong; 7, stridulatory comb, slide
mounted. Scale line 0.5mm.

Northern Territory and Western Australian water bugs

Figs 8-9. Anisops thienemanni male, front leg, inner view, slide mounted: 8, (Elkedra) front leg; 9, (War
Rock) femur. Scale line 0.5mm.

Figs 10-12. Anisops thienemanni male (Elkedra): 10, head and pronotum, dorsal view. Scale line 1mm.
11, stridulatory comb, (War Rock); 12, stridulatory comb. Scale line 0.5mm.

I. Lansbury

Figs 13-14. Anisops thienemanni male (Elkedra): 13. rostral
prong (War Rock); 14, rostral prong. Scale line 0.5mm.

from north-western Australia, Sir Graham Moore
Island, based on two females (Brooks 1951:
353), is thought to be a misidentification. The
occurrence of A. gratus north of Alice Springs is
probably nearing the extreme northern edge of
its range. During 1979, 20 habitats were sam¬
pled within 100 km of Alice Springs, A. gratus
being found only in six habitats, in numbers
consistently below 10 specimens.

Anisops thienemanni Lundblad
(Figs 8-14)

Anisops thienemanni Lundblad, 1933: 167-
168; -Brooks 1951: 413-416; -Sweeney 1965:
88-89; -Lansbury 1969: 446-448.

Material examined. NTM Entomological
collection. Northern Territory, Elkedra River,
two males, 10 females.

Description. Males 6.97-7.50 mm long, fe¬
males 7.13-7.80 mm long.

The relative size of both sexes from Elkedra
more or less fall within the parameters of Brooks
(1951) and Lansbury (1969). This species was
originally described from Java, Diengplateau.
Australian recordsextend over most of the Eyrean
and Bassian regions, and the species is common
in Tasmania, whereas A. gratus is only known
from one locality in Tasmania.

The Elkedra male front femur (Fig. 8) is
distally acuminate, the ridge on the inner sur¬
face only extending a short distance across the
femur. The femur of the more robust form from
south-western Australia (War Rock, 15 km north
of Morawa,29°05’Sl 16°00’E) is rounded distally
and the inner ridge is longer (Fig. 9). The
stridulatory combs of both forms are similar
(Figs 11-12). The head and pronotum of the
Elkedra male (Fig. 10)hasalongpronotum.The
rostral prong of the Elkedra male (Fig. 14)
differs in geometry and size when compared
with the Morawa form (Fig. 13) which is con¬
stricted towards the apex compared with the
explanate rostral prong of the Elkedra form.

Distribution. Anisops thienemanni does not
seem to have been collected from the northern
regions of the Northern Territory and north
Queensland. In some areas of southern Queens¬
land, New South Wales and elsewhere, A.
thienemanni is often found with A gratus in
saline inland waters up to 14.0 mg/1. Anisops
thienemanni is one of the first water-bugs to
colonise ephemeral pools and is frequently at¬
tracted to artificial light.

The variation between the forms of A.
thienemanni suggests that they may be a group
of sibling species and the ‘typical form’ from
Java is not conspecific with the Australian form.

Northern Territory and Western Australian water bugs

Figs 15-19. Anisops elkedraensis sp. nov. paratype male: 15, front leg (stridulatory comb omitted), slide mounted, inner view,
16, rostral prong. Scale line 0.5mm. 17, head and pronotum, dorsal view. Scale line 1 mm. 18, front tibia; 19, front tarsus, slide
mounted, inner view. Scale line 0.5mm.

Anisops nasuta Fieber

Anisops nasuta Fieber, 1851: 484-485 (see
Brooks 1951:416-418 for extensive bibliogra¬
phy and synonymy 1851-1941); -Lansbury 1969:
440-441; -Lansbury 1991: 113.

Material examined. NTM Entomological
collection. Northern Territory, Elkedra River, 3
males, 15 females. Torresian-Eyrean species,
widespread in SE Pacific region.

Anisops paracrinita Brooks

Anisops paracrinita Brooks, 1951: 329-331;
-Lansbury 1964: 57-58; -Lansbury 1969:434; -
Lansbury 1991: 111.

Material examined. NTM Entomological
collection. Northern Territory, Elkedra River, 1
male.

A Torresian species not previously recorded
from Alice Springs area.

I. Lansbury

Anisops elkedraensis sp. nov.

(Figs 15-19)

Type material. HOLOTYPE - NTM 1.551,
male. Northern Territory, Elkedra River, 21 ° 13 ’S
-135°085’E, 28 September 1982, coll. P. Homer,

H. Larson and K. Bishop. PARATYPES - 8
males, NTM 1.552-561; 8 females, NTM 1.562-
571, both lots same data as for holotype.

Description. Malcs5.39-5.80 mm long, 1.49-

I. 51 mm maximum width, females 6.20-6.80
mm long, 1.66-1.72 mm maximum width.

Structure of male. Eyes and vertex more or
less evenly rounded. Greatest width of head
0.89x pronotal humeral width, 5.9x anterior
width of vertex and 3x median head length.
Synthlipsis 0.27x anterior width of vertex. Me¬
dian head length 0.7x median pronotal length.
Pronotal humeral width 2.43x median length,
lateral margins 0.6x median length, diverging
and straight. Posterior margin deeply emarginate
(Fig. 17). Facial tubercle slightly flattened. Ba¬
sal width of labrum slightly longer than median

length. Lower margin of facial tubercle and
labrum with scattered erect short hairs (Fig. 16).
Rostral prong as in Figure 16. Chaetotaxy of
front leg as in Figure 15 (stridulatory comb
omitted). Front tibia (Fig. 18) stridulatory comb
with approximately 20 pegs. Front tarsus (Fig-
19) with five evenly spaced setae along inner
margin.

Structure of female. Greatest head width 0.87x
pronotal humeral width, 6.36x anterior width of
vertex and 3.50x median head length. Synthlipsis
0.28x anterior width of vertex. Median head
length 0.66x pronotal length, humeral width
2.60x median length. Pronotum and facial tu¬
bercle and labrum similar to male.

Colouration. Ethanol specimens. Eyes red¬
dish brown. Vertex, pronotum and elytra par¬
tially hyaline greyish brown. Legs pale yellow.
Thorax and ventrites dark brown-black, keel
and lateral margins of connexivum pale yellow.

Remarks. These specimens key out to couplet
89 in Brooks (1951), Anisops exigera Horvath,
which is a smaller species, with males 4.3-4.5

Figs 20-21. Anisops hayesi sp. nov. paratype male: 20, front leg, inner view, slide mounted. Scale line 1mm.
21, stridulatory comb. Scale line 0.5mm.

Northern Territory and Western Australian water bugs

24 -

Figs 24-25. Anisops ungarinyin sp. nov. Holotype male: 24, head and pronotum, dorsal view; 25, rostral prong. Scale line 1 mm.

I. Lansbury

mm long and having a short rostral prong. In
Lansbury (1969), A. elkedraensis keys out as A.
paraexigera Lansbury, which has a small group
of proximal setae along the inner margin of the
front tarsus; A. elkedraensis sp. nov. differs in
having a row of five prominent setae along the
front tarsus.

Etymology. The species is named for the type
area, Elkedra River.

Anisops hayesi sp. nov.

(Figs 20-23)

Type material. HOLOTYPE- male. NTM I.
584, Northern Territory, Alice Springs, John
Hayes Rock-hole near Trephina Gorge, 27 April
1979, coll. I. Lansbury. PARATYPES - 3 males,
5 females, same data as holotype, in Oxford, 1
male, 2 females, NTM 1. 585-586, same data as
for holotype, in Northern Territory Museum.

Additional material. Northern Territory,
Alice Springs, Standley Chasm, 50 km west of
Alice Springs, 16 April 1985, coll. F.S. Truxal,
1 male, in Oxford.

Description. Males 7.63 mm long, greatest
width 2.49 mm, females 7.50 mm long, greatest
width 2.49 mm.

Structure of male. Viewed dorsally.head large,
eyes voluminous and extending beyond anterior
margin of vertex (Fig. 23). Greatest head width
0.96x pronotal humeral width, 8.58x anterior
width of vertex and 3.2x head length. Synthlipsis
subequal to anterior width of vertex. Dorsal
inner lateral eye margins slightly concave. Me¬
dian head length 0.7x median pronotal length.
Pronotal humeral width 2.4x median length,
lateral margins diverging and concave. Poste¬
rior margin medially emarginate. Pronotum with
two (1+1) groups of minute pits either side of
midline. Facial tubercle raised (Fig. 22) basally
just above labrum with a dark brown pointed
tubercle. Basal labral width slightly greater than
median length with almost erect stout short
hairs down part of rostrum. Rostral prong as in
Figure 22. Chaetotaxy of front leg as in Figure
20. Stridulatory comb as in Figure 21.

Structure of female. Viewed dorsally, head
large, eyes extending beyond anterior width of
vertex. Greatest head width 0.89x pronotal hu¬
meral width, 7.14x anterior width of vertex and
3.30x median head length. Synthlipsis subequal
in width to anterior width of vertex. Dorsal inner
lateral margins of eyes almost straight. Median
head length 0.62x median pronotal length.
Pronotal humeral width 2.30x median length.

Lateral margins diverging, straight; posterior
margin almost straight. Facial tubercle slightly
raised. Basal labral width broader than median
length. Frons lined with fine long hairs extend¬
ing to facial tubercle, labrum with semi-erect
short hairs.

Coloration. Eyes black. Vertex, pronotum
and scutellum greyish with brown infuscations.
Hemelytrahyaline, appearing dark due to brown-
black dorsum showing through. Thorax ventrally
yellowish brown, ventrites shining black. Legs
yellowish brown, mid-femora with elongate dark
brown infuscations.

Remarks. This species keys out to couplet 53
in Brooks (1951), Anisops windi Brooks and A.
philippinensis Brooks; A. hayesi differs from
both species by its much larger size and the
tubercle on the lower margin of the facial tuber¬
cle. In Lansbury (1969), it keys out to couplet A.
windi and A. tasmaniaensis Brooks; the projec¬
tion on the facial tubercle distinguishes A. hayesi
from these and all other described Australian
species.

Etymology. The species is named for the type
area, John Hayes Rockhole, near Trephina Gorge.

Anisops ungarinyin sp. nov.

(Figs 24-28)

Type material. HOLOTYPE - Western Aus¬
tralian Museum (WAM)94/710, male, Western
Australia, Kimberley region, 28.7 km from
Barnett River Crossing, Phillips Gorge, 3 Feb¬
ruary 1985. Holotype, in South Australian Mu¬
seum (SAM).

Description. Structure of male. Male 6.05mm
long, maximum width 1,82mm. Viewed dorsally,
eyes large and voluminous. Greatest head width
1.03x pronotal width, 12.50x anterior width of
vertex and 3.60x median length. Synthlipsis
subequal to anterior width of vertex. Dorsal
inner lateral eye margins shallowly concave.
Median head length subequal to median pronotal
length, lateral margins straight, 0.60x median
length. Posterior margin medially shallowly
emarginate (Fig. 24). Facial tubercle not raised
(Fig. 25). Labrum short, 1.5x broader than long.
Facial tubercle and labrum with short hairs.
Rostrum long (Fig. 25). Front femur as in Figure
26, front tibia moderately spinose, stridulatory
comb with 18 pegs (Fig. 27). Front tarsus with
median row of prominent setae (Fig. 28).

Coloration. Eyes black with irregular yellow¬
ish brown blotches. Vertex, pronotum and
scutellum pale creamy yellow. Hemelytra hyaline,

Northern Territory and Western Australian water bugs

Figs 26-28. Anisops ungarinyin sp. nov. holotype male. Front leg, slide mounted, inner view: 26, femur, 27, tibia;
28, tarsus. Scale line 0.5mm.

dorsal abdominal bluish-black pigmentation
showing through. Thorax yellowish brown cen¬
trally, centrites black, keel and connexival mar¬
gins pale yellow, legs yellow.

Remarks. Anisops ungarinyin sp. nov. keys
out to couplet 55 in Brooks (1951), Anisops
doris Kirkaldy and A. tasmaniaensis. The former
has 50+ pegs in the stridulatory comb;
tasmaniaensis is larger, 7.5-7.9mm long and is
confined to Tasmania and a few localities in
Victoria and New South Wales. In Lansbury
(1969) A. ungarinyin keys out as Anisops
occipitalis Breddin, the basal rostral segment of
A. occipitalis is wider than the base of the fourth
segment, the synthlipsis is about half the ante¬
rior width of the vertex compared with the
subequal synthlipsis/anteriorwidthof the vertex
of Anisops ungarinyin sp. nov.

Etymology. The specific name ‘ ungarinyin’
refers to both the language and tribal name of the
Aboriginal people of the central Kimberley re¬
gion, Western Australia.

REFERENCES

Brooks, G.T. 1951. A revision of the genus Anisops
(Notonectidae, Hemiptera). University of Kan¬
sas Science Bulletin 34(1): 301-519.

Fieber, F.X. 1851. Rhynchotographieen.
Abhandtungen Kbniglichen Bbhmischen
Gessellschaft der Wissenschaften 5: 469-486.
Hale, H.M. 1923. Studies in Australian aquatic
Hemiptera. Records of the South Australian
Museum 11(3): 397-424.

Lansbury, I. 1964. The genus Anisops in Australia
(Hemiptera-Heteroptera, Notonectidae). Jour-

I. Lansbury

nal of the Entomological Society of Queensland
3: 52-65.

Lansbury, I. 1969. The genus Anisops in Australia
(Hemiptera-Heteroptera, Notonectidae). Jour¬
nal of Natural History 3: 433-458.

Lansbury, I. 1991. Naucoridae and Notonectidae
(Hemiptera-Heteroptera) of the Northern Terri¬
tory, Australia. The Beagle,Records of the North¬
ern Territory Museum of Arts and Sciences
8(1): 103-144.

Lundblad, O. 1933. Zur Kenntnis der aquatilen und
semi-aquatilen Hemipteren von Sumatra, Java
und Bali. Archiv JUr Hydrohiologie Supplement
12(4): 1-195, 263-489.

Sweeney, A.W. 1965. The distribution of the
Notonectidae (Hemiptcra) in south-eastern Aus¬
tralia. Proceedings of the Linnean Society of
New South Wales 90(1): 87-94.

Accepted 27 September, 1995

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:75-88

TWO NEW SPECIES OF CTENOTUS (REPTILIA, SCINCIDAE) FROM THE

NORTHERN TERRITORY.

PAUL HORNER

Museum and Art Gallery of the Northern Territory,

P.O. Box 4646, Darwin, NT 0801, Australia.

ABSTRACT

Two new species of Ctenotus Storr (Reptilia, Scincidae), C. astictus sp. nov. and C.
stuarti sp. nov. are described and their habitat preferences and distribution in the
Northern Territory are discussed. Both species are closely allied to C. arnhemensis
Storr and are distinguished from that species by a combination of body pattern, and
morphological and meristic characteristics. Ctenotus arnhemensis is redescribed.

Keywords: Reptilia, Scincidae, Ctenotus, new species, Northern Territory.

INTRODUCTION

The genus Ctenotus Storr, 1964, is the most
s Peciose taxon of scincid lizards found in Aus-
tr alia. It includes 82 species (Cogger 1992)
w hich are distributed throughout the continent
^nd only one of these extends to southern New
Guinea.

Ctenotus species are found through a diverse
array of habitats, ranging from arid deserts to
r °pical savanna woodland, with many being
tyhipatrically distributed. Syntopy is common
' v [ t h 12 species sharing the same habitat in
Uru National Park, Northern Territory (Reid
et al. 1993 ) and seven species in one region of
and south west Australia(Pianka 1969). Ctenotus
J! most diverse in the arid regions of Australia.
°wever, of the 40 species known to occur in the
°rthern Territory, 22 are found in the arid
I 9 q tre anc ^ ' ^ in the tropical north (Horner
. 2 ). Many of the northern species are special-

Se d tropical forms and it is now becoming
Pparent that these may have very restricted
ls tributions, exhibiting a high degree of habitat
• s Pecifi c j t y

The ‘fop End’ of the Northern Territory
Resents a paradox in the knowledge of its fauna.

Ccess t0 many extensive tracts of land is very
b - , r ^nd these have been sparsely surveyed
°gically, if at all. In contrast, the 19,757
4 U are kilometre expanse of centrally located
a adu National Park boasts one of the best

known faunas in Australia. Field work by the
author in certain accessible areas of the Top End
has shown that specimens putatively identified
as C. arnhemensis exhibited two forms of non-
random colour variation. Following the avail¬
ability of sufficient specimens these forms were
considered to represent new species and are here
described and compared to C. arnhemensis,
whose original description is herein expanded.
The two new species and C. arnhemensis form a
closely allied species group which is termed the
C. arnhemensis species-complex.

MATERIALS AND METHODS

A detailed morphometric and meristic analy¬
sis was made on a series of 25 specimens of two
previously undescribed species of Ctenotus and
on nine specimens of C. arnhemensis. Fourteen
scale counts and eleven measurements were
made on each specimen. Measurements were
made with micrometer adjusted callipers and/or
a steel rule. Counts of supraocular, supraciliary,
ciliary, supralabial and infralabial scales,
subdigital lamellae and ear lobules were made
on both sides of the body.

Of the measurements and counts taken, the
following require individual definition:

1. Ear-snout (head) length: measured from
the anterior margin of the ear orifice to the tip of
the rostral scale;

P. Homer

2. Eye-snout (snout) length: measured from
the anterior margin of the orbit to the tip of the
rostral scale;

3. Limb length: measured along the poste¬
rior edge, from the body wall to the tip of the
longest toe (claw excluded);

4. Fourth toe length: measured from the
angle between the third and fourth toes to the tip
of the fourth toe (claw excluded);

5. Forelimb to snout length: measured from
the anterior margin of the forelimb, at the body
wall, to the tip of die rostral scale;

6. Paravertebral scales: counted from first
scale behind parietals to a point midway be¬
tween the hindlimbs.

Scalation and body pattern nomenclature fol¬
lows that of Homer (1992).

The following abbreviations are used in the
text: AM, Australian Museum; NTM, Museum
and Art Gallery of the Northern Territory.

SYSTEMATICS

Members of the C. arnhemensis species-com¬
plex have the following character states in com¬
mon with most other Ctenotus. Body elongate,
slender, as wide as deep. Tail slender, round in
section and tapers to point. Well-developed
pentadactyl limbs with long, pointed digits.
Dorsal scales smooth with four parallel rows of
larger paravertebral scales extending from
nuchals to tail. Lateral scales small and smooth.
Scales on abdomen twice as large as lateral
scales. Subcaudal scales single and very large.
Head wider than deep. Snout pointed. Lower
eyelid moveable, opaque. Nasal scales separated
by rostral and frontonasal scales. Frontoparietal
scales paired. Interparietal scale distinct. Pari¬
etal scales large and in contact behind
interparietal scale. Two loreal scales. Upper and
lower preocular scales present, lower much the
larger. Presubocular scale present. Single pri¬
mary temporal scale and two secondary
temporals. Auricular opening large, ovoid with
obtusely pointed lobules on anterior margin.

The following five character states in combi¬
nation place them in the C. lesueurii species-
group of Storr (Storr 1981; Storr et al. 1981).
Digits slightly compressed with moderately
broad, dark calli on the subdigital lamellae. Four
supraocular scales, first three contacting fron¬
tal, second larger than first, third and fourth.
Supraciliary scales very disparate in size (fourth
to eighth much smaller than others). Ear lobules

large andgraded in size. Colour pattern includes
a wide, dark, white-edged vertebral stripe.

Members of the complex are distinguished
from other members of the C. lesueurii species-
group by the following six character states in
combination. Body dorsal surface with three
dark longitudinal stripes. Prefrontal scales not
in contact. Maximum snout-vent length less
than 60 mm. Midbody scale rows 26 or less.
Usually seven supralabials. Subdigital lamellae
under the fourth toe, 21-26.

The three members of the C. arnhemensis
species-complex are distinguished from each
other by acombination ofmorphological, meristic
and colour pattern characteristics.

Ctenotus astictus sp. nov.

(Figs 1-2)

Type material. HOLOTYPE-NTM R.l 1252,
14 kilometres west of Numbulwar on Ngukurr
road, 14°14’S 135°36’E, Northern Territory,
coll. I. Archibald, 07 June 1983. PARATYPES
- NORTHERN TERRITORY: NTM R.l 1250-
51, same data as holotype; NTM R. 14183, Lake
Eames, Vanderlin Island, Sir Edward Pellew
Group, 15°41’S 137°02’E; NTM R.16113,
R.16134-35, R.16140-44, Cadell River cross¬
ing, 12°15’S 134°26’E; NTM R.16177, Liver¬
pool River crossing, 12°22’S 134°07’E; NTM
R.19151, R. 19153, Red Point, Marchinbar Is¬
land, Wessel Islands, 11°16’S 136°35’E; NTM
R.20252, Bumaga Island, Wessel Islands,
11°46’S 136°05’E.

Additional material. Northern Territory: ju¬
veniles, NTM R. 11248-49, same data as holotype.

Diagnosis. Ctenotus astictus sp. nov. is dis¬
tinguished from C. arnhemensis and C. stuarti
sp. nov. by having a reduced, simple lateral
pattern which lacks any form of pale spots or
blotches and by having generally fewerear lobules
(mean of 2.9 vs means of 4.8 and 4.1 respec¬
tively). Additionally, the relative lengths of the
limbs and tai 1 are generally greater in C. astictus
sp. nov. than in C. arnhemensis and C. stuarti
sp. nov.

Ctenotus astictus sp. nov. is superficially
similar to well patterned specimens of C.
vertebralis, but may be distinguished by pos¬
sessing a grey-brown rather than black upper
lateral zone, wider second supraocular scale and
fewer ear lobules and supralabial scales.

Description. Head. Snout length 39-49%
(mean = 42.4%) ofhead length. Prefrontal scales
usually separated by frontal scale (87%), occa-

New skinks from the Northern Territory

Fig. 1 . Paratype of Crenotusastictus sp. nov. (NTM R. 16143) from Cadell River crossing, Arnhem Land, Northern Territory,
photographed in life.

sionally in narrow contact. Nuchal scale pairs
usually four (69%), occasionally five (25%) or
three (6%). Supraciliary scales usually ten (81%),
occasionally nine or eleven, fourth to eighth
markedly smaller than others. Ciliary scales
usually ten (44%) or eleven (44%), occasionally
nine or twelve. Supralabial scales usually seven
(94%) occasionally eight, fifth under orbit.
Infralabial scales usually seven (93%) occasion¬
ally eight, twocontacting postmental. Ear lobules
usually three (69%) or four (25%), rarely two
( 6 %).

Body. Snout-vent length to 51.9 mm (mean =
47.1 mm). Axilla to groin length 49-58% (mean
= 52.3%) of snout-vent length. Tail length 211-
239% (mean = 228.3%) of snout-vent length.
Paravertebral scales 48-56 (mean = 52.7).
Midbody scale rows usually 24 (87%), occasion¬
ally 26 (12%).

Limbs. Forelimb length 25-32% (mean =
28.4%) of snout-vent length. Hindlimb length
42-52% (mean = 48.2%) of snout-vent length.
Fourth toe length 15-19% (mean = 17.3%) of
snout-vent length. Subdigital lamellae under
fourth toe 21-25 (mean - 22.4) excluding claw.

Colour and pattern (in spirit). The basic
body pattern is of a reduced form, consisting of
a simple arrangement of smooth-edged, longitu¬
dinal stripes and zones of ground colour. Pale
spotting is virtually absent.

Body. Dorsal surface of body mid-brown with
simple pattern of smooth edged stripes. Dark black¬
ish-brown vertebral stripe, half as wide as para¬
vertebral scales, extends from anteriornuchal scales
to base of tail. Vertebral stripe margined by white
paravertebral stripes which are about one third as
wide as vertebral stripe. Paravertebral stripes bor¬
dered by zone of mid-brown background colour,
about as wide as paravertebral scales, which ex¬
tends from parietal scales onto tail. Outer margins
of background colour zones bordered by prominent
blackish-brown laterodorsal stripes, about three
quarters as wide as vertebral stripe, which extend
from outer edge of parietal scales onto tail. Outer
margins of laterodorsal stripes bordered by distinct
white dorsolateral stripes, about half as wide as
vertebral stripe, which extend from outer edge of
the fourth supraciliary scales onto tail.

Lateral surface of body grey-brown patterned
with two pale, smooth edged stripes. Broad grey-

P. Homer

brown upper lateral zone extends from above
auricular opening onto tail. This is separated from
mid-lateral zone by prominent white upper lat¬
eral stripe, about one third as wide as upper
lateral zone, which extends from upper posterior
margin of auricular opening, above the limbs, onto
tail. Grey-brown mid-lateral zone, about three
quarters as wide as upper lateral zone, extends
from posterior margin of auricular opening to
base of tail. This is separated from lower lateral
zone by an obscure, pale lower lateral stripe,
about as wide as upper lateral stripe, which ex¬
tends from lower posterior margin of auricular
opening to forelimb, behind which it continues to
hindlimb. Obscure, grey-brown lowerlateral zone,
about half as wide as upper lateral zone, extends
from below auricular opening to hindlimb, and
coalesces into immaculate whitish ventral surface.

Head. Snout light brown. Dark mottling on
frontal, frontoparietal, interparietal and parietal
scales. Temporal region mid-brown, patterned
with broken pale stripe, (or series of pale
blotches), which extends from anterior temporal
scales to above auricular opening. White
subocular stripe extends from lower preocular
scale to upper margin of auricular opening.
Supralabials light brown with pale margins be¬
tween scales. Infralabials mottled brown and
off-white. White ventral surface changing to
cream on chin.

Limbs. Mid-brown background colouration
on dorsal surface of forelimbs separating three
black stripes. Cream on ventral surface.
Hindlimbs similar but with four black stripes.

Tail. Light brown on dorsal surface. Basal
portion has remnants of dark vertebral stripe.
Continuations of dark laterodorsal stripes, pale
dorsolateral stripes, upper lateral zone and pale
upper lateral stripe extend along length of tail.
Cream on ventral surface.

Details of holotype. (NTM R.l 1252). Snout-
vent length 50.3 mm; tail length 106.0 mm;
axilla to groin length 29.3 mm; forelimb length
13.7 mm; hindlimb length 21.4 mm; fourth toe
length 7.8 mm; head width 5.4 mm; head depth
4.9 mm; eye to snout length 4.2 mm; ear to snout
length 9.9 mm; forelimb to snout length 16.9
mm; nasals separated; prefrontals separated;
supraciliaries ten on both sides; ciliaries ten on
both sides; supralabials seven on both sides;
infralabials seven on both sides; nuchals four
pairs; ear lobules two on left side, three on right
side, uppermost largest; subdigital lamellae of
fourth toe 22 on both sides; midbody scale rows
24; paravertebral scales 54.

Variation. The 16 specimens analysed are
generally uniform in most characteristics of
scalation, morphology, colouration and pattern.
Any variation in morphology and meristics is
summarised in Table 1.

Fig. 2. Holotype of Ctenotus astictus sp. nov. (NTM R.11252). Head scalation in a) right lateral and b) dorsal view.
Scale bar - 5 mm.

New skinks from the Northern Territory

Distribution. The known distribution of C.
astictus sp. nov. is north-eastern Northern Ter¬
ritory. It has been collected near Numbulwar
and at the Cadell and Liverpool river crossings
in Arnhem Land, and on Marchinbar and
Bumaga Islands in the Wessel Islands group and
Vanderlin Island in the Sir Edward Pellew Group.

Habitat. Mainland specimens were found in
low-lying, white sandy flats vegetated by open
Eucalyptus woodlands or tall, open Acacia
shrubland. All were collected from sites with a
ground storey of perennial grass tussocks on a
pale sandy substrate, adjacent to watercourses.
The Vanderlin Island site supported Eucalyptus
-dominated low, open woodland with a open,
hummock-grassland understorey of Plectrachne
pungens. Specimens from the Wessel Islands
were collected at three sites, descriptions of
which are: 1, flat bordering rocky coastline with
a mixed strand community of scattered tall
Casuarina equisitifolia, small dense patches of
strandline, coastal monsoon vine forest and
mixed low tussock grasses and forbs, on loamy
humic sand substrate; 2, gently undulating, ex¬
tensive sand flat behind beach with a low wood¬
land of Acacia plectocarpa and Melaleuca ner¬
vosa, and a sparse understorey of small shrubs
and Triodia microstachya; 3, sandy crest and
upper slopes of low hind-dune with a mosaic of
small, dense thickets of coastal monsoon vine
forest, and a low, mid-dense understorey of
herbs and tussock grasses plus scattered Triodia
microstachya hummocks.

Ctenotus astictus sp. nov. specimens have
been recorded using shallow ground burrows for
shelter and small juveniles (26-29 mm in snout-
vent length) were collected near Numbulwar in
June.

Specimens of C. essingtonii were also col¬
lected at the Cadell and Liverpool river sites.

Etymology. The species name astictus is de¬
rived from the Greek term stiktos which means
‘spotted’. It refers to the unspotted nature of the
body pattern in comparison to C. arnhemensis
and C. stuarti sp. nov.

Ctenotus stuarti sp. nov.

(Figs 3-4)

Type material. HOLOTYPE -NTM R. 13723,
Swim Creek, Opium Creek Station, 12°34’30"S
131 °49 ’ 30"E, Northern Territory, coll. P. Homer,
24 April 1988. PARATYPES - NORTHERN
TERRITORY: NTM R. 13086-87, same data as
holotype except 24 April 1985; NTM R. 13722,

R. 13724-27, same data as holotype; NTM
R. 17453, Kapalga, 12° 40’S 132° 22’E.

Diagnosis. Ctenotus stuarti sp. nov. is dis¬
tinguished from C. astictus sp. nov. by having
a complex lateral pattern of stripes, pale spots
and blotches, and by having generally more ear
lobules (mean of 4.1 vs mean of 2.9). Addition¬
ally, the relative lengths of the limbs are gener¬
ally less in C. stuarti sp. nov. than in C. astictus
sp. nov.. Ctenotus stuarti sp. nov. is distin¬
guished from C. arnhemensis by having a gen¬
erally lower number of subdigital lamellae un¬
der the fourth toe (mean of 20.9 vs mean of
22.9) and by having generally more nuchal
scale pairs (mean of 4.7 vs mean of 3.8). Also,
the head of C. stuarti sp. nov. is usually deeper
than that of C. arnhemensis (head depth as
percentage of head width, mean of 88.6% vs
mean of 85.7%). In addition, the continuous,
smooth-edged, white dorsolateral stripes and
dark laterodorsal stripes found in C.
arnhemensis and other members of the C.
lesueurii species-group are broken into ragged-
edged, semi-continuous series of flecks and
streaks in C. stuarti sp. nov.

Description. Head. Snout length 38.6-54.2%
of head length. Prefrontal scales usually sepa¬
rated by frontal scale (89%), occasionally in
narrow contact. Nuchal scale pairs usually five
(89%), occasionally four. Supraciliary scales
usually nine (63%), occasionally ten, fourth to
eighth markedly smaller than others. Ciliary
scales usually eleven (50%) or ten (38%), occa¬
sionally twelve. Usually seven supralabial scales
(88%), occasionally eight, fifth under orbit.
Infralabial scales seven, two contacting
postmental. Ear lobules usually four (78%),
occasionally with five or six.

Body. Snout-vent length to 52.0 mm (mean =
48.8 mm). Axilla to groin length 49-54% (mean
= 51.6%) of snout-vent length. Tail length 204-
226% (mean = 215%) of snout-vent length.
Paravertebral scales 50-58 (mean = 54.3).
Midbody scale rows usually 24 (44%) or 26
(33%), occasionally 25.

Limbs. Forelimb length 24-28% (mean=26%)
of snout-vent length. Hindlimb length 41-45%
(mean = 43.5%) of snout-vent length. Fourth toe
length 13-16% (mean = 14.4%) of snout-vent
length. Subdigital lamellae under fourth toe 18-
23 (mean = 20.9) excluding claw.

Colour and pattern (in spirit). The basic
body pattern consists of a complex arrangement
of longitudinal stripes, zones of ground colour,
pale spots and blotches.

P. Homer

Fig. 3. a) Holotype of Ctenotus stuarti sp. nov. (NTM R. 13723) from Swim Creek, Opium Creek Station, Northern Territory
photographed in life, b) Detail of forebody. Note discontinuous series of streaks representing dark laterodorsal and pale
dorsolateral lines.

New skinks from the Northern Territory

Body. Dorsal surface olive-brown with com¬
plex pattern of smooth and ragged edged stripes.
Dark, smooth-edged, blackish-brown vertebral
stripe, half as wide as paravertebral scales, ex¬
tends from posterior edge of parietal scales to
base of tail. Vertebral stripe margined by smooth-
edged, white paravertebral stripes about one
third as wide as vertebral stripe. Paravertebral
stripes bordered by zone of olive-brown back¬
ground colour, about as wide as paravertebral
scales, which extends from parietal scales onto
tail. Outer margins of background colour zones
bordered by obscure, narrow, laterodorsal series
of irregular blackish-brown flecks and streaks,
which extend, as an irregular, discontinuous
stripe, from upper edge of primary temporal
scales onto tail. This dark laterodorsal series
merges with distinct discontinuous, white,
dorsolateral stripe. Formed by ragged-edged,
series of flecks and streaks, this continuously
broken stripe, about half as wide as vertebral
stripe, extends from outer edge of fourth
supraciliary scales onto tail. Interspaces in the
series often as wide as white streaks, coloured
with mingling of blackish-brown laterodorsal
series and brown upper lateral zone.

Lateral surface of body brown, patterned with
pale spots and blotches and, at least posteriorly,
a pale mid-lateral stripe. Broad brown upper
lateral zone extends from above auricular open¬

ing onto tail, patterned with one or two irregular
series of 18-22 small pale blotches; zone sepa¬
rated from lower lateral zone by white mid¬
lateral stripe, about one third as wide as upper
lateral zone, which extends from above forelimb
to hindlimb, behind which it continues onto tail.
This stripe is often indistinct and may be broken
on anterior half of body, as series of elongated
blotches. Between forelimb and auricular open¬
ing, mid-lateral stripe represented by two or
three pale blotches. Mottled grey-white lower
lateral zone, about half as wide as upper lateral
zone, extends from below auricular opening to
hindlimb, patterned anteriorly by numerous,
obscure, pale spots and blotches; and coalesces
into immaculate whitish ventral surface.

Head. Snout light brown. Dark mottling on
supraoculars, frontal, frontoparietal,
interparietal, parietal and nuchal scales. Tem¬
poral region patterned with three or four irregu¬
lar pale blotches. Ragged white subocular stripe
extends from second loreal scale to mid-upper
margin of subocular scale (fifth supralabial),
then extends along upper margin of posterior
supralabial scales as irregular series of three or
four pale blotches. Supralabials mottled grey-
white with pale margins between scales giving
barred appearance. Infralabials mottled white-
grey. White ventral surface changing to cream
on chin.

Fig. 4. Holotype of Ctenotus sluarli sp. nov. (NTM R.13723). Head scalation in a) right lateral and b) dorsal view.
Scale bar - 5 mm.

P. Homer

Limbs. Mid-brown background colouration
on dorsal surface of forelimbs separating three
black stripes. Cream on ventral surface.
Hindlimbs similar but with four black stripes.

Tail. Light brown on dorsal surface. Basal
portion with remnants of dark vertebral stripe.
Continuations of dark laterodorsal stripes, pale
dorsolateral stripes and pale mid-lateral stripes
extend along length of tail. Cream on ventral
surface.

Details of holotype. (NTM R. 13723). Snout-
vent length 52.0 mm; tail length 106.0 mm;
axilla to groin length 26.1 mm; forelimb length
13.3 mm; hindlimb length 21.3 mm; fourth toe
length 7.4 mm; head width 6.5 mm; head depth
5.8 mm; eye to snout length 4.3 mm; ear to snout
length 10.5 mm; forelimb to snout length 18.0
mm; nasals separated; prefrontals separated;
supraciliaries ten on both sides; ciliaries eleven
on both sides; supralabials seven on both sides;
infralabials seven on both sides; nuchals five
pairs; ear lobules four on both sides, second from
top largest; subdigital lamellae of fourth toe 20
on both sides; midbody scale rows 26;
paravertebral scales 53.

Heart and liver tissue samples were taken
from the holotype at death. These are lodged in
the South Australian Museum’s tissue bank
under the number SAM-EBU H73.

Variation. The nine specimens analysed are
generally uniform in most characteristics of
scalation, morphology, colouration and pattern.
Any variation in morphology and meristics is
summarised in Table 1. One individual from
Kapalga differs slightly by possessing more
continuous laterodorsal and dorsolateral stripes.
In this specimen they are only occasionally
interrupted by an intrusion of background col¬
our.

Distribution. The known distribution of C.
stuarti sp. nov. is restricted to two localities.
Opium Creek Station, 100 kilometres east of
Darwin, where it was collected adjacent to Swim
Creek (this property is a recent subdivision of
Point Stuart Station), and CSIRO Research Sta¬
tion, Kapalga, approximately 65 kilometres fur¬
ther east.

Habitat. Specimens from Swim Creek were
collected on a gently sloping, white sandy flat
supporting a low open woodland. The site is low
lying, adjoining a shallow, seasonal drainage
channel, extending about 100 metres, which
empties into perennial Swim Creek. The site’s
dominant vegetation types are low, scattered
tree and shrub species and a moderately thick to

sparse groundstorey of perennial grass and sedge
tussocks. Surrounded by open Eucalyptus forest
on other soil types. Prominent shrub/tree species
are Jacksonia dilatata, Melaleuca cajuputi,
Verticordia cunninghamii, Calytrix extipulata
and Grevillea pteridifolia. Numerous C. stuarti
sp. nov. specimens were seen in the immediate
area and observations on their activity indicate
that the species retreats to shallow ground bur¬
rows for protection, occasional ly climbs into low
vegetation whilst foraging and (during the
monsoonal wet season) remains in areas where
the substrate is fully saturated and covered by a
centimetre or two of water. Two other Ctenotus
species were recorded from the site, C. essingtonii
and C. slorri.

Specimens from Kapalga were collected in
open (wet) woodland on deep sandy soil, near a
seasonal seepage zone.

Etymology. The new species is named for
John McDouall Stuart, first explorer to cross the
continent from south to north and survive. He
reached the north coast, in the vicinity of the
type locality, in July 1862. The name also al¬
ludes to Point Stuart, a natural feature of the
Northern Territory coast located close to the
type locality.

Ctenotus arnhemensis Storr, 1981
(Figs 5-6)

Ctenotus arnhemensis Storr, 1981: 127,
Jabiluka, Northern Territory.

Material examined. HOLOTYPE - AM
R.88613, Jabiluka Mineral Lease, Kakadu Na¬
tional Park, Northern Territory, 12°33’S
132°55’E. PARATYPES - NTM R.7620-23,
same data as holotype; NTM R.20253, same
data as holotype, except 12°34.6’S 132°54.5’E;
NTM R.20254, same data as holotype, except
12°30.8’S 132°54.0’E; NTM R.20255-56, same
data as holotype.

Diagnosis. Ctenotus arnhemensis is distin¬
guished from C. astictus sp. nov. by having a
complex lateral pattern of stripes, pale spots and
blotches and by having generally more ear lobules
(mean of 4.8 vs mean of 2.9). Additionally, the
relative length of the limbs are usually less in C.
arnhemensis than in C. astictus sp. nov. Ctenotus
arnhemensis is distinguished from C. stuarti sp.
nov. by having a generally higher number of
subdigital lamellae under the fourth toe (mean
of 22.9 vs mean of 21.0) and by having generally
fewer nuchal scale pairs (mean of 3.8 vs mean of
4.7). Also, the head of C. arnhemensis is usually

New skinks from the Northern Territory

Fig. 5. Ctenotus arnhemensis (NTM R.20256) from Jabiluka, Northern Territory, photographed in life.

less deep than that of C. stuarti sp. nov. (head
depth as percentage of head width, mean of
85.7% vs mean of 88.6%). In addition, C.
arnhemensis has continuous, smooth-edged,
white dorsolateral stripes and dark laterodorsal
stripes, features which in C. stuarti sp. nov. are
broken into ragged-edged, semi-continuous se¬
ries of flecks and streaks.

Description. Figures in squared brackets in¬
dicate measurements from the original descrip¬
tion differing from those of this analysis:

Head. Snout length 39-48% (mean = 41.3%) of
head length. Prefrontal scales separated by fron¬
tal scale. Nuchal scale pairs usually four (89%),
occasionally three [3-5, mean=3.6]. Supraciliary
scales usually ten (54%) or nine (46%), fourth to
eighth markedly smaller than others. Ciliary
scales usually eleven (56%) or ten (33%), occa¬
sionally nine [10-14, mean = 11.5]. Usually
seven supralabial scales (90%), occasionally
eight, fifth under orbit. Infralabial scales usually
seven (89%), occasionally eight, two contacting
postmental. Ear lobules usually five (67%), oc¬
casionally four or six [5-7, mean = 5.5].

Body. Snout-vent length to 55.1 mm (mean =
51.1 mm). Axilla to groin length 49-58% (mean
= 53.9%) of snout-vent length. Tail length 196-
244% (mean=218%) of snout-vent length [191-
237%, mean = 207%]. Paravertebral scales 50-
59 (mean - 53.9). Midbody scale rows usually
24 (78%), occasionally 26 [23-27, mean = 24.8].

Limbs. Forelimb length 23-28% (mean =
25.8%) of snout-vent length [24-29%, mean =
25.4%]. Hindlimb length 41-49% (mean =
44.2%) of snout-vent length [40-48%, mean =
43%]. Fourth toe length 13-18%(mean= 14.8%)
of snout-vent length. Subdigital lamellae under
fourth toe 21-26 (mean = 22.9) excluding claw
[20-23, mean = 21.4].

Colour and pattern (in spirit). The basic
body pattern consists of a complex arrangement
of smooth-edged, longitudinal stripes, zones of
ground colour, pale spots and blotches.

Body. Dorsal surface mid-brown, patterned
with smooth edged stripes. Dark, blackish-brown
vertebral stripe, half as wide as paravertebral
scales, extends from anterior nuchal scales to
base of tail. Vertebral stripe margined by white

P. Homer

paravertebral stripes which are about one third
as wide as vertebral stripe. Paravertebral stripes
bordered by zones of mid-brown background
colour, about as wide as paravertebral scales,
which extend from parietal scales onto tail.
Outer margins of background colour zones bor¬
dered by prominent, blackish-brown laterodorsal
stripes, about three quarters as wide as vertebral
stripe, which extend from outer edge of parietal
scales onto tail. Outer margins of laterodorsal
stripes bordered by distinct, white dorsolateral
stripes, about half as wide as vertebral stripe,
which extend from outer edge of fourth
supraciliary scales onto tail.

Lateral surface of body brown, patterned with
pale spots and blotches and pale mid-lateral
stripe. Broad blackish-brown upper lateral zone
extends from above auricular opening onto tail
and patterned with single, regular series of 19-
21 large pale blotches. Prominent white mid¬
lateral stripe, about one third as wide as upper
lateral zone, extends from upper posterior mar¬
gin of auricular opening, above limbs onto tail.
Between forelimb and auricular opening, mid¬
lateral stripe may be broken into series of two or
three pale blotches. Brown lower lateral zone,
about half as wide as upper lateral zone, extends
from below auricular opening to hindlimb. This
is patterned anteriorly by four or five pale
blotches. Some specimens also with obscure,
ragged, pale lower lateral stripe which may only

be visible posteriorly. Where present, this coa¬
lesces into immaculate whitish ventral surface.

Head. Snout light brown. Dark mottling on
frontal, frontoparietal, interparietal and parietal
scales. Temporal region brown, patterned with
three or four irregular pale blotches. White
subocular stripe extends from first loreal scale to
mid-upper margin of sixth supralabial scale.
Supralabials light brown with pale margins be¬
tween the scales. Infralabials mottled brown and
off-white. White ventral surface changing to
cream on chin.

Limbs. Mid-brown background colouration
on dorsal surface of forelimbs separating three
black stripes. Cream on ventral surface.
Hindlimbs similar but with four black stripes.

Tail. Light brown on dorsal surface. Basal
portion has remnants of dark vertebral stripe.
Continuations of dark laterodorsal stripes, pale
dorsolateral stripes, upper lateral zone and pale
mid-lateral stripes extend along length of tail.
Cream on ventral surface.

Distribution. The known distribution of C.
amhemensis is extremely limited, being re¬
stricted to within, or the near vicinity of, the
Jabiluka and Ranger Project Areas located within
Stage One of Kakadu National Park.

Habitat. R. Sadlier (pers. comm.) collected
the type series on white sandy flats adjacent to
the Jabiluka sandstone outlier. Vegetation at
these sites consisted of open woodland with a

Fig. 6. Holotype of Ctenotus arnhemensis (AM R.88613). Head
Scale bar =■ 5 mm.

New skinks from the Northern Territory

Fig. 7. Map of the Top End region of the Northern Territory showing the distribution of Ctenotus arnhemensis (triangles),
C. astictus sp. nov. (circles) and C. stuani sp. nov. (squares). Outlined area indicates position of Kakadu National ar .

Discriminant function 1

Fig. 8. Plot of discriminant analysis on six variables recorded from 34 specimens of Ctenotus arnhemensis (sp. 1), C. astictus
sp. nov. (sp.2) and C. stuarti sp. nov. (sp.3).

P. Homer

sparse middle storey of shrubs (eg. Grevillea
spp.) and a groundstorey of sedge and grass
tussocks. Part of the type series was also col¬
lected from tall grassland, on humic silt substrate,
at the northern end of nearby Island Billabong.
Two other Ctenotus species were sympatric at
these sites, C. essingtonii and C. storri.

Variation. The nine specimens analysed are
generally uniform in most morphometric and
meristic characters, as well as in body colouration
and pattern. Any variation in morphology and
meristics is summarised in Table 1.

DISCUSSION

As an aid to identification of Ctenotus in
Western Australia, Storr (Storr 1981; Storr et
al. 1981) distributed the species from that re¬
gion among ten species-groups. Based on char¬

acters in common, these species-groups are not
necessarily natural but are useful in clustering
similar species together. Wilson and Knowles
(1988) tentatively recognised twelve species-
groups which include all Ctenotus species.
Ctenotus arnhemensis, C. astictus and C. stuarti
have character states in common that place
them in Storr’s C. lesueurii species-group, the
content of which now includes: C. arcanus
Czechura and Wombey, G. arnhemensis Storr,
C. astictus, C. borealis Homer and King, C.
brachyonyx Storr, C. capricorni Storr, C.
coggeri Sadlier, C. eurydice Czechura and
Wombey, C. eutaenius Storr, C. fallens Storr,
C. helenae Storr, C. hypatia Ingram and
Czechura, C. ingrami Czechura and Wombey,
C. inornatus (Gray), C. lateralis Storr, C.
lesueurii (Dumeril and Bibron), C. mastigura
Storr, C. monticola Storr, C. nullum Ingram
and Czechura, C. robustus Storr, C. saxatilis

Table 1. Morphometric and meristic characteristics of Ctenotus arnhemensis , C. astictus sp. nov. and C. stuarti sp. nov..
Measurements are in mm. Tail lengths were measured from original tails only. (SD - standard deviation).

Characteristics

Ctenotus arnhemensis

n-9

mean SD range

Ctenotus astictus sp. nov.
n - 16

mean SD range

Ctenotus stuarti sp. nov.
n-9

mean SD range

Snout-vent length (SVL)

51.11

3.49

45.9-55.1

47.12

3.22

40.8-51.9

48.85

2.90

42.1-52.0

Tail length

109.33

10.93

94.0-122.1

109.44

8.11

96.0-124.0

111.50

7.78

106.0-117.0

Axilla-groin length

27.63

3.26

32.2-31.6

24.65

2.05

20.0-29.3

25.23

1.90

21.0-27.8

Forelimb length

13.13

1.35

10.8-14.9

13.34

0.69

122-14.6

12.74

0.56

11.7-13.3

Hindi imb length

22.50

0.71

21.5-23.3

22.64

1.36

19.9-25.1

21.24

1.22

18.9-23.1

Fourth toe length

7.55

0.36

7.1-8.2

8.12

0.49

7.1-9.1

7.01

0.48

6.4-7.9

Head width

6.14

0.35

5.^63

5.65

0.54

4.6-63

5.78

0.41

5.2-63

Head depth

5.27

0.50

42-5.8

4.92

0.47

3.9-5.9

5.12

0.45

4.4-5.8

Eye-snout (snout) length

4.13

0.07

40-4.2

4.07

0.46

3.2-5.1

4.15

0.59

3.4-5.4

Ear-snout (head) length

10.02

0.61

8.7-10.8

9.61

0.76

7.7-10.6

9.87

0.58

8.8-103

Fore limb-snout length

16.83

0.53

15.6-17.5

16.71

1.26

13.8-18.6

16.47

1.24

14.7-18.1

No. of supraciliaiy scales

9.56

0.46

9-10

9.97

0.22

9-11

9.37

0.52

9-10

No. of ciliary scales

10.28

0.62

9-11

10.47

0.72

9-12

10.62

0.69

10-12

No. of supralabial scales

7.05

0.17

7-8

7.03

0.12

7-8

7.06

0.18

7-8

No. of infralabial scales

7.11

0.33

7-8

7.03

0.13

7-8

7.00

0.00

No. of nuchal scale pairs

3.78

0.36

3-4

4.03

0.46

3-5

4.67

0.35

4-5

No. of ear lobules

4.78

0.62

2.94

0.44

2-4

4.11

0.60

3-6

No. of 4th toe lamellae

22.89

1.36

21-26

22.37

1.13

21-25

20.89

1.34

18-23

No. of midbody scale rows

24.44

0.88

2426

24.25

0.68

24-26

24.89

0.93

24-26

No. of paravertebral scales

53.89

3.02

50-59

52.75

1.95

48-56

54.33

2.91

50-58

ratios

Tail length to SVL

217.98

16.37

196.3-243.9

228.29

8.28

210.6-238.9

214.83

15.53

203.8-225.8

Axilla-groin length to SVL

53.92

3.15

49.458.1

52.30

2.01

49.1-58.2

51.62

1.68

49.3-53.7

Fonelimb length to SVL

25.84

1.32

23.5-27.7

28.38

1.73

24.7-323

26.12

1.08

24.4-27.8

Hindlimb length to SVL

44.16

2.53

41.5-49.0

48.17

2.42

42.6-51.8

43.50

1.42

41.1-45.1

4th toe length to SVL

14.84

1.50

133-17.5

17.26

0.87

15.5-18.6

14.37

0.96

12.8-16.1

Head depth to head width

85.73

5.69

74.6-91.5

87.23

6.09

75.0-96.3

88.60

4.90

80.6-96.6

Orbit-snout to Ear-snout

41.33

2.74

38.9-47.8

42.38

2.61

39.5-49.4

42.06

5.04

38.6-54.2

Ear-snout to SVL

19.63

0.94

183-21.0

20.37

0.91

18.6-21.9

20.23

0.44

19.5-20.9

Forelimb-snout to SVL

33.05

2.34

29.9-37.1

35.48

1.54

33.4-373

33.75

1.63

31.2-35.7

New skinks from the Northern Territory

Storr, C. severus Storr, C. spaldingii (Macleay),
C. stuarti, C. taeniolatus (White, ex Shaw), C.
terrareginae Ingram and Czechura, and C.
vertebral is Rankin and Gillam.

Ctenotus amhemensis, C. astictus and C.
stuarti have no single morphometric or meristic
character to distinguish them. Their body pat¬
terns are distinctive, but require support to war¬
rant species separation. Table 1 shows that many
variables recorded for each of these species have
different means, but have considerable overlap.
Six of these, those used to distinguish between
the three taxa, were subjected to a Discriminant
Analysis using the program Statgraphics (ver¬
sion 5). Figure 8 illustrates that this analysis
separated individual specimens into three dis¬
crete groups, corresponding to C. amhemensis
(sp.l), C. astictus (sp.2) and C. stuarti (sp.3).
With high canonical correlation coefficients
(above 0.78) and low significance levels (less
than 0.0001) indicating that the discriminant
functions are significant, the discrimination
analysis results show that several characters,
used in combination, can distinguish between
these three species.

The three species appear to be allopatrically
distributed (Fig. 7). Kakadu National Park
(19,757 km 2 ) bisects the Top End of the North¬
ern Territory and has been the subject of numer¬
ous, intensive faunal surveys, none of which
have recorded the presence of C. astictus or have
found C. amhemensis other than in, or adjacent
to, the Jabiluka and Ranger Project Areas (200
km 2 ). Ctenotus stuarti has been recorded from
Kakadu National Park, at Kapalga in Stage Two
but is apparently absent from Stages One and
Three. Interestingly, Kapalga has been the sub¬
ject of faunal studies for many years, but C.
stuarti is only known from two individuals
collected at a single study site. Thus, the three
species appear to have restricted distributions, a
finding that is supported by the number of other
Ctenotus species in the Top End only known
from small geographic areas. Examples of low¬
land forms of these are: C. borealis Homer and
King; C. gagudju Sadlier, Wombey and
Braithwaite; C. hilli Storr; C. kurnbudj Sadlier,
Wombey and Braithwaite; and, C. storri Rankin
(see distributions in Homer 1992).

Habitat preferences of C. amhemensis, C.
astictus and C. stuarti are similar, all occurring
on white, sandy flats with an open woodland or
open shrubland upperstorey, and a groundstorey
of perennial grass and/or sedge tussocks. Each

species also shares sympatry with C. essingtonii
and, in the case of C. amhemensis and C. stuarti,
C. storri. Sadlier, Wombey and Braithwaite
(1986) discussed the habitat preferences of
Ctenotus species in the Alligator Rivers region
of the Top End. Of eight species, they recorded
three examples of two species occurring in eco¬
logical sympatry (syntopy) and one example of
three species being syntopic (C. amhemensis, C.
essingtonii and C. storri). They explained the
region’s diversity of Ctenotus as being due to the
diverse habitats found in the region, and diver¬
sity within habitats by the size differences be¬
tween species and in the way that similar-sized
species partition resources, such as microhabitat,
food and activity period.

The addition of two new species of Ctenotus
to the fauna of the Top End means that, in the
Northern Territory, tropical forms now almost
rival in numberthoseof the arid centre. With the
strong possibility of more species to be described
from the region, the monsoonal north of Aus¬
tralia may, in future, rival the arid centre in
diversity of Ctenotus.

ACKNOWLEDGEMENTS

For their contributions of specimens I am
grateful to I. Archibald, A. Fisher, J. Woinarski
and C. Trainor. R. Sadlier (AM) facilitated the
loan of type material in his care and provided
habitat details for sites in the Jabiluka Mineral
Lease. For the detailed description of Wessel
Islands sites I am especially grateful to A. Fisher.

REFERENCES

Cogger, H.G. 1992. Reptiles and Amphibians of
Australia. Fifth Edition. Reed Books: Chatswood.
Homer, P. 1992. Skinks of the Northern Territory.
Handbook Series No.2. Northern Territory Mu¬
seum of Arts and Sciences: Darwin.

Pianka, E.R. 1969. Sympatry of desert lizards
(Ctenotus ) in Western Australia. Ecology 50:
1012 - 1030.

Reid, J.R.W., Kerle, J.A., Baker, L. and Jones, K.R.
1993. Reptiles and frogs. In: Reid, J.R.W., Kerle,
J.A. and Morton, S.R. (eds). Uluru Fauna. The
distribution and abundance of vertebrate fauna
of Uluru (Ayers Rock - Mount Olga) National
Park, N.T. Kowari 4: 58-68.

Sadlier, R.A., Wombey, J.C. and Braithwaite, R.W.
1986. Ctenotus kurnbudj and Ctenotus gagudju,
two new lizards (Scincidae) from the Alligator

P. Homer

Rivers region of the Northern Territory. The
Beagle, Occasional Papers of the Northern Ter¬
ritory Museum of Arts and Sciences 2: 95-103.

Storr, G.M. 1964. Ctenotus, a new generic name for
a group of Australian skinks. Western Austral¬
ian Naturalist 9: 84-85.

Storr, G.M. 1981. Ten new Ctenotus (Lacertilia:
Scincidae) from Australia. Records of the West¬
ern Australian Museum 9: 125-146.

Storr, G.M., Smith, L.A. and Johnstone, R.E. 1981.
Lizards of Western Australia. 1. Skinks. Univer¬
sity of Western Australia Press and Western
Australian Museum: Perth.

Wilson, S.K. and Knowles, D.G. 1988. Australia's
reptiles. A photographic reference to the terres¬
trial reptiles of Australia. William Collins Pty
Ltd: Sydney.

Accepted: 29 August, 1994

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:89-99

DENTATE AND RELATED STONE BIFACE POINTS FROM
NORTHERN AUSTRALIA.

KIM AKERMAN 1 AND PETER BINDON 2
' Museum and Art Gallery of the Northern Territory,

P.O. Box 4646, Darwin, NT 0801, Australia.

2 Western Australian Museum,

Francis St, Perth, WA 6000, Australia.

ABSTRACT

The Kimberley spear point as a typological entity has always been problematical. A
brief discussion on some types of ethnographic and prehistoric biface points from the
Northern Territory and the Kimberley region of Western Australia seeks to clarity
some of these problems. A hidierto unrecognised, but distinct, form of stone biface
point with dentate margins from the Kimberley is described, and a clarification of
terms used for margin treatment of stone points is presented.

Keywords: stone points, biface, pressure flaking, margin treatment, Kimberley,
northern Australia.

INTRODUCTION

In this paper we define terms used to describe
the edge treatment of bifacially pressure-flaked
stone points, and using these definitions we
present a classification of some types of these
artefacts found in northern and north western
Australia. A previously undescribed point is
illustrated and discussed.

To date, little regional and typological analy¬
sis of bifacially flaked point industries has been
undertaken. Previous work has focussed mainly
on possible differentiation of types among sym¬
metrical unifacial points (Campbell and Noone
1943;Campbell 1960:509-524; Mulvaney 1975:
319-221; McCarthy 1976: 42). The most note¬
worthy exceptions are the analysis of the points
from Yarar shelter. Port Keats, undertaken by
Flood (1970), Schrire’ s (1982) analysis of exca¬
vated material from five sites in the Alligator
Rivers region of West Arnhem Land, and Allen
and Barton’s (1989) analysis and discussion of
points from Ngarradj Warde Djobkeng in the
same area. ( Fig.l).

Flood (1970) analysed a point series exca¬
vated from Yarar rock shelter in the Northern
Territory, separating them initially into
unifacially trimmed and bifacially trimmed

groups. Dortch (1977) reviewed the northern
Australian occurrence of various unifacially
flaked points and included a type he referred to
as ‘Kimberley backed points’( Dortch 1977:

117). Here, we concentrate on bifacially flaked
points, especially those which are invasively
flaked using pressure techniques.

Three terms are used to describe specialized
margin treatment that may occur on pressure
flaked points:

dentate, describes a margin with more-or-less
regularly spaced projections or teeth separated
by notches that are wider than the teeth;

denticulate, describes regularly spaced pro¬
jections which are separated by notches that are
of similar width or narrower than the teeth
themselves; and

serrated, which refers to extremely small or
fine projections usually triangular in outline and
separated from each otherby equally fine notches.

Throughout the Kimberley Aboriginal lan¬
guage groups, the terms used for these three
types of projection can be glossed as “teeth”. To
produce each “tooth” on a dentate margin, the
knapper must remove a series of flakes from the
embayment between the projections. These small
flakes are removed from both faces of the arte¬
fact and produce scars which are long and

K. Akerman and P. Bindon

shallow. The finished margin thus presents, in
plan, a series of rectangular gaps which separate
sub-rectangular projections. Flakes forming this
edge are most invasive towards the longitudinal
axis of the point, and the abrupt or steeper sides
of each projecting tooth are shaped by crushing
the brittle margins that remain after removal of
the invasive flakes.

Projections on a denticulate margin are rela¬
tively smaller than those on the dentate margin,
even though the completed artefacts may be of
similar size and proportion. Usually the pres¬
sure flakes which produce the embayment be¬
tween the projections are removed bifacially
from the margin and more or less perpendicular
to the central axis of the point.

Serrations usually protrude less than 2 mm
from the body of the artefact. As a result of the
overlapping flake scars produced when notch¬
ing, serrations are often triangular in plan with
pointed apices.

NORTHERN AUSTRALIAN BIFACE
POINTS

Two basic but distinctive forms of bifacially
flaked points were made in the ethnographic
present in northern Australia, and two other
distinctive types are seemingly prehistoric. The
two ethnographic points are the Wanji Biface
(Wanji Point) and the Kimberley Point, while the

two prehistoric point types have yet to be named
in the literature. For convenience, the first pre¬
historic point type we describe will be called the
Northern Territory Triangular Point, the second
prehistoric point type is described in detail and
given the name Kimberley Dentate Point.

The Wanji Biface. In western Arnhem Land
and adjacent areas to the west and south-west,
the Wanji Biface (McCarthy 1976:44) was made
until very recently. This point is made by percus¬
sion flaking of fissile materials such as indurated
slate or other rocks possessing cleavage planes
(Fig. 2). Large glass points (> 14 cm in length)
made from plate glass or from the flat sides of
square faced spirits bottles are associated with
Wanji Points and apparently are a post-contact
manifestation of the stone prototypes. These
glass examples do not exhibit invasive pressure
flaking. The chipping, which may be done either
by percussion or pressure and removes only
short flakes, has been applied to provide plan
symmetry to the artefact, but margins are rarely
as acute as the margins of points from the
Kimberley. When found hafted, these glass points
are invariably mounted directly into single¬
piece bamboo shafts with beeswax cement and a
fibre cord binding, rather than being resin
mounted on the composite shafts typical of the
Kimberley spears. We suggest that these glass
points are derived from the stone Wanji Points
and are not related to the pressure flaked ser¬
rated glass points of the Kimberley region.

Biface points from northern Australia

Fig. 2. Wanji Points. A, Pine Creek; B, Arnhem Land (after McCarthy 1976: Fig. 14(No. 1)); C, Arnhem Land (after Dahl
1927: Plate 4 (No. 6)); D, Oenpelli, Arnhem Land (after McCarthy 1960: Plate 12 (No. 5)).

Northern Territory Triangular Points. This
distinctive prehistoric biface point type occurs
in south western Arnhem Land and adjacent
areas. In plan, this point resembles an isosceles
triangle with a width to length ratio of approxi¬
mately 1:3. The base appears either straight or
slightly curved and the remaining two margins
are more or less straight (Fig. 3). These points
commonly range from 40 - 90 mm in length.
They are produced by carefully controlled, deli¬
cate percussion flaking, with minimal pressure
flaking being undertaken to straighten the mar¬
gins and form the tip. These points possess a
more-or-less constant relationship between form
and mass not observable in the other types of
bifacial points discussed here. These points ap¬
pear, however, to be prehistoric, there being no
known ethnographic record or hafted examples
existing.

Kimberley Points. The third form of point is
manufactured usingpressure technique on blanks
prepared by percussion. Stone, glass and ce¬
ramic points of this type are found in the Kim¬
berley and the area immediately to the east. It is
this type of point which is usually referred to as
the “Kimberley point”.

Fig. 3. Prehistoric biface points, central northern and north¬
western Northern Territory.

K. Akerman and P. Bindon

Previously, the term ‘Kimberley point’ has
been applied to what we consider to be a range
of points, usually flaked bifacially, but not al¬
ways by pressure flaking (McCarthy 1976: 42).
Although some authors (White and O’Connell
1982: 122) restrict the term to those points bear¬
ing serrated edges, we consider that even this
usage is unsatisfactory. There are two basic prob¬
lems with the term Kimberley point as it has
been used in earlier literature. Firstly, the name
has been applied historically to more than one
type of artefact. Within the Kimberley region
itself, at least four types of bifacially flaked points
can be distinguished on the basis of their margin
treatments. Typologically it is unsatisfactory to
describe these points with one all-embracing

term. Secondly, although pressure-flaked bifae e
points are common on Kimberley open sites and
in excavations, there is no evidence as yet that
pressure flaking of stone points originated in the
Kimberley. As noted below, we believe that the
likely dissemination centre for pressure flaking
technology may lie much further east.

We suggest that the term Kimberley Point
should only be used as a general term for biface
points that are manufactured by pressure flak-
ing. Pressure flaking proceeds from prepared
platforms and each successive series of flakes is
taken along one margin and refines half of each
face alternately. The shape of Kimberley Points
varies from ovate to lanceolate, with rounded
bases and acuminate tips. Margins are invari-

Fig. 4. Denticulate and serrated pressure flaked stone Kimberley Points. Top row: denticulate points. A, Lissadell Station; B,
Halls Creek; C, N ingbing Station; D, Christmas Creek Station; E, Lamboo Station. Bottom row: outlinesof serrated stone points
from Kunmunya Mission.

Biface points from northern Australia

ably either denticulate or serrate (Fig. 4).

Museum collections in Australia hold many
examples of Kimberley Points derived from both
ethnographic and archaeological sources. Broad
examples produced from sections of glass bot¬
tles usually have relatively short scalar retouch
on the concave face, while on narrow points the
concavity may be obliterated. All known ethno¬
graphic examples have serrated and/or
denticulate lateral margins. Some glass serrated
points (Fig. 5) may have 6-8 teeth per 10 mm of
margin, while denticulate stone points possess
2-3 teeth per 10 mm. In the Kimberley today,
serration and denticulation of points does not
imply or reflect any cultural differentiation,
either spatially or temporally. Our observations
of contemporary point manufacture demonstrate
that both types of margin treatment may be
utilised by any one Aboriginal knapper, and the
output of a single craftsman usually includes
points with both forms of margin treatment. No
typological distinctions are drawn by contempo¬
rary Aboriginals between points with serrated or
denticulated margins.

Love (1936: 74-75), Elkin (1948: 110-113)
and Tindale (1985: 1-33) provided detailed de-

Fig. 5. Pressure flaked glass points with serrated margins,
Kimberley, Western Australia. The point at right has the fine
tip protected by cord wrapping.

scriptions of the manufacture of what are com¬
monly called Kimberley Points. Such tools are
illustrated in Figure 6. Pressure flaking of stone
or glass points in the Kimberley today has be¬
come a lost art, although there are still a few
people alive who knapped up to the mid 1980s
but who are now aged and infirm. In addition,
there are many Aboriginal people who, while
not having practised the craft, have observed
knappers in action in the recent past and can
often provide information on the topic of lithic
technology.

A broad range of raw materials were exploited
in the Kimberley for the production of ethno¬
graphic biface points. While some of these ma¬
terials are widely available, others are more
restricted in their distribution. Historically at
Kunmunya, small (15-30 mm long) finely ser¬
rated points were generally made of translucent
orange, white, yellow and red agates and
chalcedonies that occur as small nodules in the
basalt derived soils. Larger points from this area
were made from silcretes and the opaque green
to black cherts that occur throughout the Kim¬
berley plateau. In the west Kimberley, high-
grade quartz crystal was often used in the north¬
ern portions of the Napier Range and fossiliferous
chert used in the southern parts. Variegated
cherts were commonly used for points along the
Ord Valley and the area immediately east of die
Ord River. In the southern and south-eastern
Kimberley, extensive outcrops of white chert
were exploited as raw material. Across the Kim¬
berley plateau and in the Ord River Basin,
cobbles weathered from conglomerates provided
a source of high-grade silcrete. To the east at
Timber Creek, a distinctive grey merging to
pink chert, derived from the Bardia Chert Mem¬
ber of the Skull Creek Formation, was used as
raw material for the manufacture of both
unifacially and bifacially flaked points. No doubt,
with intensification of archaeological research
in the north, further discrete and identifiable
sources of raw material will be located.

Kimberley Dentate Points. There is how¬
ever, a class of biface points diat is quite differ¬
ent from those described above. Rather than
being lanceolate in form with a rounded base
and a short acuminate tip typical of the most
recently made Kimberley biface points, these
points are generally very narrow in relation to
their length (Fig. 7). Adjacent to the rounded
base and proceeding toward the pointed tip, the
margin bears a series of large irregular teeth

K. Akerman and P. Bindon

separated by wide deep notches. Notching may
extend for about half the overall length of the
artefact, which then tapers more or less smoothly
into a long drawn out tip. Teeth on opposite sides
of each margin may be roughly aligned so that a
degree of bilateral symmetry is maintained. As
these points only occur in the Kimberley, it is
proposed that they be called Kimberley Dentate
Points.

Kimberley Dentate Points vary considerably
in size. The largest examined is 115 mm long,
with the majority ranging between 30 mm and
50 mm in length. Smaller points may have been
used as projectile points, but the apparent fragil¬
ity of larger examples suggests their use as
prestige trade goods. The width/thickness ratio
across the teeth is approximately 3.0, but only
about 2.0 between the wide and deep notches.
The length/width ratio varies between 4.0 and
5.0. Many of these points we have examined
have lost the tip and the projecting teeth of
others are damaged. Larger points that have
apparently lost the tip, probably through acci¬
dent rather than use, appear to have been subse¬
quently rejuvenated by pressure flaking. Rather
than creating a new elongate tip, which would
require the removal of a substantial mass of
material the distal edges have been brought to a
simple ogival tip. Such a rejuvenated point is
illustrated by Noone (1943: 244, Fig. 4).The
toothed portion of the margins occupies ap¬
proximately 0.3 to 0.6 of the overall length of the
point. On the large points, the length of the
dentate margin is proportionally greater than
that area of a smaller point. The narrow parallel¬
sided tips are rhomboidal in section and rela¬
tively shorter in the larger points of this type
than in the smaller examples.

Two manufacturing techniques were used to
produce these points, depending on the size and
shape of the available blanks. Large points were
usually made on either thin tabular pieces or on
large flakes of a range of silicified sedimentary
rocks and tuffs. Some examples exhibit the
remains of a primary ventral surface indicating
that the blank was originally a flake. These
pieces or flakes were reduced to preforms by
percussion flaking. This reduced their width
and thickness and ensured that a straight longi¬
tudinal profile was obtained. Pressure flaking
was then used to notch the proximal margins to
a depth of up to 6 mm, leaving the teeth between
the notches standing clear. The notches may be

up to 9 mm wide at the base and bear scars
indicating that multiple flakes were removed
from each face in the notching process. The
projecting teeth are usually irregular in width
and shape and create an overall illusion of
complexity. In some instances the outer margins
of the teeth are serrated. The inner side of each
notch is roughly aligned with that of each adja¬
cent notch and the teeth are set in balanced pairs
on opposite margins so the greater mass of the
artefact retains an elegant bilateral symmetry.
Immediately in front of the most distal pair of
notches, the point contracts relatively abruptly
before extending to a long spike-like tip. The
margins of the zone of contraction may be roughly
denticulated while the tip bears even smooth
edges.

The point is refined, prior to notching, by
collateral pressure flaking undertaken on an
anvil, and the invasive flakes removed are rela¬
tively short (7.0 mm). At the proximal section of
the point, the pressure flaking docs not obliter¬
ate the preforming percussion flake scars. At the
narrower distal end and at the tip, the pressure
flake scars meet at the midline creating the
rhomboidal-sectioned tip.

Smaller Kimberley Dentate Points (< 50 mm
long) are generally made directly on narrow,
pointed flake-blades. The usual raw material is
a white chert, common in the southern Kimber¬
ley; silicified tuff and a green, medium-grained
chert may also be utilised. Although these points
are bifacially worked, there is often no attempt
to straighten the profile and the points may
retain the curved longitudinal profile of the
original flake-blade. The use of resin to haft
spear points in the Kimberley allows even mark¬
edly curved points to be hafted with the axis of
the point aligned with the shaft, minimising any
effect the curvature would have either on the
flight or penetration performance of the spear
(Akerman 1978). Preforming by percussion is
unusual and the notching and shaping of the
proximal section may involve removal of a sin¬
gle series of flakes removed from both faces
along each margin. Constriction of the tip re¬
quires the removal of several series of collateral
flakes from each margin and face. Smaller points,
made by first percussion preforming pieces of
suitable material, resemble the larger examples
except, as with all the smaller points, the length
of the tip is proportionally longer in relation to
the length of the dentate margin. The base of all

Biface points from northern Australia

Fig. 6. Bark wallet and pressure flaking tools. Kunmunya Mission. A, cord wrapped bark wallet containing worked stone points
andpressure flaking tools. Worora name buru. ru (Western Australian Museum (WAM 10093); B, hardwood indenter. Worora
name karindjalp: C. kangaroo ulna indenter. Worora same ljurmba (WAM 10102); D, kangaroo fibula indenter, Worora name
tingkalja. (WAM 10107); E-F, wire indenters.

K. Akerman and P. Bindon

examples, whether large or small, is rounded
and reduced to the width of the body between the
bases of the most proximal pair of notches.

For the Kimberley Dentate, form or shape
appears to be determined on a cultural rather
than a technological basis. It must however be
recognised that by making these points on nar¬
row flake-blades, the need for extensive pres¬
sure-flaking was minimised and there may have
been recognised savings in both time and effort
by prehistoric knappers at least in regard to the
manufacture of the smaller dentate points. Gen¬
erally, with regard to variation in the dimen¬
sions of Australian biface points, the raw mate¬
rial used or the technique of manufacture may,
in some instances, determine the size range of
points produced. The small agate points from
the Kunmunya area and the slate Wanji biface
points respectively, reflect such constraints.

Occurrence and distribution of Kimberley
Dentate Points. While no points of this type
have yet been recognised in a situation that
would allow them to be placed into a chronologi¬
cal sequence, they do appear to have a definite
spatial distribution in the Kimberley, being rela¬
tively common on open sites, particularly in the
southern part of the Fitzroy River drainage
basin. These open sites are of considerable inter¬
est, as the artefacts scattered upon them appear

to be derived from industries generally regarded
as discrete entities, reflecting cultural differen¬
tiation. Tula adze-flakes, core tools, edge-ground
adzes and hatchet heads, pirri gravers, blades,
unifacial and bifacial points as well as grinding
and pounding stones and worked baler shell are
all present at many of these sites. Such an
aggregation is indicative of intense cultural
interchange, with amalgamation of elements of
northern, coastal and desertcultural suites. This
combination forms a larger and more diversified
industry than formerly existedeithertothe north
or the south. The distribution of the edge-ground
adze in the Kimberley as recorded by Akerman
and Bindon (1984: 359) corresponds closely
with that of the smaller dentate biface points.

To date, all the larger Kimberley Dentate
Points have been isolated surface finds collected
north of the Fitzroy River in the Napier Range,
the Leopold Range and at Tablelands and Ku-
runjie Stations. Smaller examples (less than 50
mm in length) are only occasionally found north
of the Fitzroy River. Apart from those observed
in the Leopold Range, it appears that many of
these larger points are traded items, an interpre¬
tation that is reinforced by their large size. This
suggests that they may have served the same
function as the contemporary large pressure-
flaked points that are known ethnographically

Fig. 7. Kimberley Dentate Points. A, Napier Downs, West Kimberley; B, Tablelands Station, East Kimberley; C, after Noone
1943: 244; D, Tablelands Station, East Kimberley; E, Bell Creek, West Kimberley. C and E have had the tips reworked.

Biface points from northern Australia

Fig. 8. Pressure flaked point with denticulate and serrated
margins (after King 1827: 68).

to represent prestige goods (Davidson 1935:
179-181). However, two points seen on a site
adjacent to Bell Creek in the Leopold Range may
have been manufactured locally. Both were made
from blades of fine-grained green chert that was
extensively exploited in this area for the manu¬
facture of points and other implements. Freshly
flaked surfaces of this chert exhibit a satin-
lustre, whilst surfaces that have been exposed to
the elements have a greasy or enamelled lustre
that could be interpreted as evidence of heat
treatment of the original material. However,
examination of a fresh surface on a broken point
revealed a satiny lustre, indicating that the glossy
lustre of the surface of the artefact was a weath¬
ering phenomenon, and not the result of heat
treatment.

Heat treatment of raw materials to enhance
their flaking qualities seems only to have been
practised in the southern portion of the area in
which Kimberley Dentate Points are found.
Consequently it is associated with the smaller

examples, although not all of these have been
subjected to this process. It should be noted,
however, that although people of the north and
central Kimberley who have made bifacially
flaked spear points in the recent past talk about
“cooking stones”, they are usually referring to
quarrying activities (Akerman 1979: 144-51).
We have neither observed true heat treatment
being practised nor found archaeological evi¬
dence for its occurrence in these northern areas.
Information on heat treatment practices in this
area can generally be regarded as unreliable and
the techniques described by contemporary Abo¬
riginal peoples are impractical, being more likely
to destroy the raw material than enhance its
flaking qualities. Historically, at least, it ap¬
pears that heat treatment to modify and improve
the flaking quality of raw materials was prac¬
tised regularly only in the south and southeast
Kimberley region.

DATING NORTHERN AUSTRALIAN
POINT TECHNOLOGIES

Jones (1985: 296) suggested that the stone
point technology in northern Australia origi¬
nated in the Alligator Rivers region of west
Arnhem Land between 5.7 and 6.2 kyrbefore the
present. In the west Kimberley, invasively flaked
bifacial points appear about 4.5 kyr BP
(O’Connor 1990:255) and 3.0kyrBP in the east
Kimberley (Dortch 1977:110). As O Connor
(1990: 208) points out, these dates “highlight
the need for larger samples before we can be
confident that our regional chronologies are
firm”. Bowdler and O’Connor (1991: 53-62)
review the literature on dates of northern point
industries and conclude that no sustainable date
earlierthan 4.5 kyrBPcan be demonstrated. The
manufacture and use of invasively flaked points
persisted until relatively recently in the Kimber¬
ley, and in historic times the techniques of point
manufacture began to diffuse eastwards
(Davidson 1935: 1701-72).

In northern parts of the Northern Territory,
judging from the ethnographic evidence, point
industries based on the manufacture of elon¬
gated flakes and pointed blades supersede
bifacially flaked point industries in the recent
past. Hafted examples of the latter type, apart
from Wanji Bifaces, are rare in museum collec¬
tions except those from the western periphery,
around about the lower Victoria River basin,

K. Akerman and P. Bindon

where an overlap in point technologies appears
to have persisted until recently. Our own obser¬
vations of point numbers on surface sites suggest
that, in the past, there was a richer point industry
in this region of overlap, as evidenced by the
presence of greater proportions ofboth bifacially
and unifacially flaked points within the stone
artefact assemblages, than can be observed in
those assemblages scattered on open sites lo¬
cated in the Kimberley. In this latter area,
bifacially flaked point technology replaced a
pointed flake/blade technology and was then
refined to a degree not found in the Northern
Territory. McCarthy (1976: 44) and White and
O’Connell (1982: 112) suggest that this refine¬
ment is a post-contact response to an external
demand for curios in combination with the avail¬
ability of new, easily worked, vitreous raw ma¬
terials such as glass and porcelain. However, it
is clear from the literature that Kimberley biface
points at contact were as refined as those pro¬
duced at later dates. King (1827: 68) illustrates
a denticulate biface point approximately 150
mm long, that is as skilfully and as regularly
flaked as any point of stone or glass manufac¬
tured subsequent to contact (Fig. 8). A photo¬
graph of this point appears in the British Mu¬
seum Handbook to the Ethnographic Collec¬
tions (1910: plate V (a)); although the extreme
tip is now missing, the very regular dentate and
denticulated margins and the flake scars con¬
form with those seen in King’s sketch. As
Etheridge (1890: 63-4) notes, this point has
“more or less square-headed teeth, themselves at
times serrated, and separated by interspaces
equal to themselves in breadth” rather than the
sharply serrated margins that he observed on
points collected in the East Kimberley.

CONCLUSION

Bifacially pressure-flaked points are still pro¬
duced occasionally within the Aboriginal com¬
munities in the Kimberley. Familiarity with the
technology of their production is now confined
to a dwindling few practitioners. Modem points
are usually leaf-shaped, made of glass, and
exhibit denticulate or serrated edges. These are
rarely, if ever, used as spear armatures. Instead
they function as trade goods and souvenirs,
much as, we suggest, did the larger Kimberley
Dentate Point in earlier times. To determine
their suitability as cultural and temporal mark¬
ers, the various kinds of bifacially flaked points

occurring in the Kimberley must be recovered in
many more stratigraphically controlled and dated
situations. Until then, these descriptions and the
typology presented should be seen as a first step
in separating and documenting some of the
various production techniques and the resultant
artefacts in the general class of tools known as
‘points’.

REFERENCES

Allen, H. and Barton, G. 1989. Ngarradj Warde
Djobkeng. Oceania Monograph 37.

Akerman, K. 1978. Notes on the Kimberley stone-
tipped spear, focussing on the hafting mecha¬
nism. Mankind 11(4): 486-489.

Akerman. K. 1979. Heat and lithic technology in the
Kimberleys W.A. Archaeology and Physical
Anthropology in Oceania 14(2): 144-151.
Akerman, K. and Bindon, P. 1984. The edge-ground
stone adze and modem counterparts in the Kim¬
berley region. Western Australia. Records of the
Western Australian Museum 11(4): 357-373.
Bowdler, S. and O’Connor, S. 1991. The dating of the
Australian small tool tradition, with new evi¬
dence from the Kimberley, W.A. Australian
Aboriginal Studies 1: 53-62.

British Museum, 1910. Handbook of the ethno¬
graphical collections. Trustees of the British
Museum: London.

Campbell, T.D. 1960. The Pirri - an interesting
Australian Aboriginal implement. Records of
the South Australian Museum 13(4): 509-524.
Campbell, T.D., and Noonc H.V.V. 1943. South Aus¬
tralian microlithic stone implements. Records of
the South Australian Museum 7(3): 281-307.
Dahl, K. 1927. In savage Australia. Houghton Mifflin:
New York.

Davidson, D.S. 1935. Archaeological problems of
northern Australia. Journal of the Royal Anthro¬
pological Institute 65: 145-184.

Dortch, C.E., 1977. Early and Late stone industrial
phases in Western Australia. In: Wright R.V.S.
(ed) Stone tools as cultural markers. Pp 104-
132. Australian Institute of Aboriginal Studies:
Canberra.

Elkin, AP. 1948. Pressure flaking in the northern
Kimberley, Australia. Man 130: 110-113.
Etheridge, R. 1890. On some beautifully formed
stone spearheads from the Kimberley, north¬
west Australia. Records of the New South Wales
Geological Survey 11: 61-65.

Flood, J.M. 1970. A point assemblage from the
Northern Territory. Archaeology and Physical
Anthropology in Oceania 5: 27-52.

Jones, R. 1985. Archaeological conclusions. In: Jones,
R. (ed.) Archaeological research in Kakadu

Biface points from northern Australia

National Park. Pp 291-298. Australian National
Parks and Wildlife Service Special Publication
013: Canberra.

King, P.P. 1827. Narrative of a survey of the coasts
of Australia, 1818-1822. London. (Australian
Facsimile Edition No. 30, 1969: Adelaide.)

Love, J.R.B. 1936. Stone-age bushmen of today.
Blackie: London.

McCarthy, F.D. 1960. The archaeology of Arnhem
Land. In: Mountford, C.P. (ed.) Records of the
American-Australian scientific expedition to
Arnhem Land. Vol 2. Melbourne University
Press: Australia.

McCarthy, FD. 1976. Australian Aboriginal stone
implements. Australian Museum Trust: Sydney.

Mulvaney, D.J. 1975. The prehistory of Australia.
Penguin: Australia.

Noone, H.V.V. 1943. Some Aboriginal stone imple¬
ments of Western Australia. Records of the
South Australian Museum 7(3): 271-280.

O’Connor, S. 1990. 30,000 years in the Kimberley: a
prehistory of the islands of the Buccaneer Archi¬
pelago and adjacent mainland, west Kimberley,
Western Australia. Unpublished PhD. thesis.
University of Western Australia.

Schrire C. 1982. The Alligator Rivers. Prehistory and
ecology in western Arnhem Land. Terra Australis
(7): Australian National University.

Tindale N.B. 1985. Australian Aboriginal techniques
of pressure-flaking stone implements. In: Plew,
M.G., Woods, J.C. and Pavesic M.G. Stone tool
analysis. Essays in honour of Don E. Crabtree.
Pp 1-33. University of New Mexico Press.

White, J.P. and O’Connell J. 1982. A prehistory of
Australia, New Guinea and Sahul. Academic
Press: Sydney.

Accepted 15 June, 1995

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:101-144

THE PALAEMONOID SHRIMP FAUNA (CRUSTACEA: DECAPODA:
CARIDEA) OF THE COBOURG PENINSULA, NORTHERN TERRITORY.

A.J. BRUCE AND K.E. COOMBES

Museum and Art Gallery of the Northern Territory,
P.O Box 4646, Darwin, 0801, Australia.

ABSTRACT

A 1981 - 1986 survey of the marine fauna of Port Essington, Cobourg Peninsula,
Arnhem Land, northern Australia, confirms the presence of 61 species of palaemonoid
shrimp. Three new species, Periclimenaeus serrula sp. nov„ Periclimenaeus solitus
sp. nov., and Periclimenes cobourgi sp. nov., are described, and a further 25 species
added to the fauna of the Northern Territory, of which Periclimenaeus stylirostris,
Periclimenes novaecaledoniae, and Periclimenes sinensis are new records for Aus¬
tralia. This report provides the first general account of the palaemonoid fauna of a
larger focal area from northern Australia, and one third of the pontoniine shrimps
known to occur in Australian waters are now known to occur in the Port Essington
region. The survey has, in total, provided eight species new to science and 34 species
have been added to the fauna of the Northern Territory.

Keywords: Faunistic survey, northern Australia, Crustacea, Decapoda, Palaemonoidea,
new species, Periclimenaeus serrula sp. nov., Periclimenaeus solitus sp. nov.,
Periclimenes cobourgi sp. nov.

INTRODUCTION

Until recent years, the tropical marine fauna
of northern Australia has attracted little scien¬
tific attention, and the caridean shrimp fauna
has remained largely undocumented. H.M.S.
A/ertpassed through PortEssington in 1882, but
did not report any shrimps from the collections
made from Arnhem Land.

From 1981 - 1986 the Northern Territory
Museum carried out a series of field studies of
the fauna of Port Essington and adjacent Cobourg
Peninsula localities. The operations were based
upon the FV Alegrias and carried out with the
financial support of the Heritage Commission.
Abundant material was obtained forstudy, which
is now preserved in the collections of the North¬
ern Territory Museum (NTM). Most of the area
studied is now included in the Gurig National
Park, established in 1981.

Much of the material collected was new to the
northern Australian fauna and some species new
to science. Some of these species have been de¬
scribed or reported on in earlier publications
(Bruce 1983a, 1983b, 1987a, 1987b, 1990a). An

account is now presented of the palaemonoid
fauna as a whole, which clearly indicates a rich
species diversity, although one that would cer¬
tainly be augmented by further collections. This
report includes descriptions of two new species
of Periclimenaeus, one new species of Peri¬
climenes, and the first records of Periclimenaeus
stylirostris, Periclimenes novaecaledoniae and
Periclimenes sinensis from Australian waters.
The area consists largely of shallow - water muddy
substrates, with rocky headlands separating sandy
bays, with areas of mangrove inshore. Coral reefs
are poorly represented, except for a well devel¬
oped reef around New Year Island and a small reef
at Coral Bay. Most of the material was hand-
collected, by hand net, using snorkel or SCUBA,
often with the use ofrotenone, with a few collec¬
tions made by a small trawl. As much material
was obtained from dive-bags containing a number
of potential host organisms, some accidental
transfers from host animals may have occurred.

Details of the stations from which sped mens were
collected are provided in Table 1 and the positions
of the localities are illustrated in Figures 1-2. Re¬
stricted synonymies only are provided, with the

101

A.J. Bruce and K.E. Coombes

original description, major name changes and
Northern Territory reports included. CL refers to the
postorbital carapace length. LWSrefers tolow water
spring tide level and LW to low water tide level.
NTM refers to the Northern Territory Museum.

SYSTEMATIC ACCOUNT

Palaemonidae Rafinesque, 1815
Subfamily Palaemoninae Rafinesque, 1815

Palaemon semmelinkii (De Man)

Leander semmelinkii De Man, 1881:137.
Palaemon (P oleander) semmelinkii - Holthuis
1950: 57-60, fig. 11.

Palaemon semmelinkii - Bruce 1987b: 58, fig. 2.

Material examined. 5 spms (1 ovig.tj), stn
CP/49, Caiman Creek, LWS, 13 May 1983, coll.
N.L. Bruce, NTM Cr.009468.

Australian distribution. Northern Territory:
Groote Eylandt (Bruce 1987b).

Further distribution. Type locality:
Makassar, Sulawesi, Indonesia. Also known
from India; Burma; the Nicobar Islands; Thai¬
land; Malaya; Singapore; Indonesia, Sumatra,
Java; and the Philippines.

Palaemon serrifer (Stimpson)

Leander serrifer Stimpson, 1860: 41; - Kemp
1925: 36.

Palaemon serrifer - Rathbun 1902: 52; -
Holthuis 1950:83-86,fig. 18;-Bruce 1987a: 57,
fig. 1; 1988b: 227.

Table 1. Summary of station data: Cobourg Peninsula.

Stn

Locality

Position

Date

Habitat

Depth

CP/2

Kennedy Bay

11°12.8'S 132°05.6’E

22.6.81

coral reef

CP/5

Coral Bay

11°11.2’S 132°03.6'E

23.6.81

coral reef

2-3m

CP/6

Coral Bay

11°11.2'S 132°03.6’E

23.6.81

off beach

3-4m

CP/7

Black Point

11°09.0'S 131°51.4'E

23.6.81

sandy beach

0.5m

CP/8

Coral Bay

11°11.2'S 132°03.6'E

24.6.81

coral reef

2-3m

CP/9

Coral Bay

11°11.3’S 132°03.75‘E

24.6.81

large shore pool

0.5m

CP/10

Black Point

11°09.0'S 132°08.2'E

18.7.81

rocky reef with algae

l-2m

CP/12

Coral Bay

1I 0 1I.3'S 132°03.75'E

19.7.81

large shore pool

< 0.5m

CP/13

Coral Bay

11°11.3'S 132°03.75’E

20.7.81

large shore pool

0.5m

CP/14

Burford Island

11°29.3*S 131°57.5'E

13.10.81

reef flat

LWS

CP/16

Wanaray Point, Trepang Bay

1I°08.0'S 131°57.7’E

14.10.81

coral reef

CP/17

Wanaray Point. Trepang Bay

i i°07.o - s i3i°58.6'E

15.10.81

coral reef

LWS

CP/18

Midjari Point. Trepang Bay

11°10.7'S 131°57.2'E

16.10.81

muddy reef flat pools

LWS

CP/20

Walford Point, Coral Bay

11°10.2'S I32°04.0'E

17.10.81

coral reef bank

LWS

CP/21

Coral Bay

U o 10.4’S I32°03.0’E

18.10.81

coral reef on rocky outcrop

l-3m

CP/23

Walford Point

11“09.4'S 132°04.0’E

19.10.81

HL81-22

NW of Orontes Reef

10°58.0’S I32°10.0'E

19.10.81

prawn trawl (3 hr)

27m

CP/24

Smith Point

11 “07.0'S I32“08.0’E

19.10.81

4-5m

CP/26

Sandy Island No. 2

11°05.5’S 132°17.0’E

20.10.81

coral reef slope

CP/27

Sandy Island No. 2

11“05.5'S 132°17.0'E

21.10.81

10m

CP/28

Sandy Island No.2

11 “07.0'S 132“17.5'E

22.10.81

coarse sandy bottom

6-7m

CP/29

Sandy Island No. 2

11°05.6'S 132°19.0'E

22.10.81

sand with rock outcrops

8-9m

CP/30

Black Point

11-09.0'S 132°08.5’E

29.4.82

rocky outcrop

10-12m

CP/33

Port Bremer

11°08.5'S 132“18.8’E

1.5.82

coral reef with Turbinaria

CP/34

Danger Point

11°39.0'S I32°20.4’E

1.5.82

sargassum bed

CP/35

Danger Point

11 “39.0'S 132°20.4'E

1.5.82

sandy beach

CP/36

Sandy Island No. 2

11°36.8'S 132°17.4'E

2.5.82

13m

CP/37

Table Head

11“13.5'S 132°10.5'E

3.5.82

rocky reef with corals

l-3m

CP/38

Table Head

11°13.5'S 132°11.5'E

4.5.82

rocky reef with Turbinaria

2-4m

CP/39

Caiman Creek

11°14.0'S 132“12.0'E

4.5.82

sandy flats & pools

CP/40

Orontes Reef

11°03.6'S 132°05.0'E

5.5.82

coral bank, offshore

NY/1

New Year Island

10°54.5'S 133°01.0'E

13.10.82

coral reef

10m

HL82-60

New Year Island

10“54.5’S 133°01.0'E

13.10.82

night light at anchor

surface

HL82-73

Oxley Island

U“10.0'S 131°58.0'E

14.10.82

night light at anchor

surface

NY/2

New Year Island

10°54.5'S 133°01.0'E

14.10.82

coral reef, few algae

10m

NY/3

New Year Island

10°55.0'S 133°OI.8’E

14.10.82

coral reef slope

16m

NY/4

McCluer Island

11 “02.0'S 132°58.5’E

16.10.82

coral reef

NY/5

McCluer Island

11°02.6’S 132°58.5’E

16.10.82

pool behind rubble ridge

NY/8

Oxley Island

10°59.0'S 132°48.8'E

19.10.32

coral reef

LWS

NY/9

Oxley Island

10°59.0'S 132°48.8’E

20.10.82

coral reef, edge of flat

LWS

102

The palaemonoid shrimp fauna of the Cobourg Peninsula

Material examined. I 9 , la", stn CP/9, Coral
Bay, 0.5 m, 24 June 1981, coll. A.J. Bruce, NTM
Cr.008541. Id", I 9 ,1 ovig. 9 ,stnCP/ 12 ,Coral Bay,
0.1-0.5 m, 19 July 1981, coll. A.J. Bruce, NTM
Cr.000909.1 juv., stn CP/26, Sandy Island No. 2,
7 m, 20 October 1981, coll. P.N. Alderslade,
J.N.A. Hooper, NTM Cr.008578. 12 spms (2
ovig. 9 ), stn CP/45, Table Head, LW, 11 May 1983,
coll. AJ. Bruce, NTM Cr.009350. 3 spms (1
ovig. 9 ), stn CP/87, Victoria Settlement, LW, 8
August 1986, coll. R.C. Willan, NTM Cr.004096.

Australian distribution. Northern Territory:
Port Essington (Bruce 1987a), Darwin Harbour
(Bruce 1988b). Queensland: Michaelmas Reef,
Capre Cay, Swain Reefs.

Further distribution. Type localities: Hong
Kong; O-Shima, Japan. Also from Southeast
Asia, from India to Vladivostok; Japan; Indone-

Tablc 1 (cont). Summary of station data: Cobourg Peninsula.

Stn Locality Position

NY/10 Oxley Island

CP/44 Table Head

CP/45 Table Head

CP/46 Table Head

CP/49 Caiman Creek
CP/51 Table Head

CP/58 Caiman Creek
CP/60 Coral Bay

CP/61 Coral Bay

CP/62 Walford Point

CP/64 Walford Point

CP/65 Middle Bay

CP/68 Coral Bay

CP/69 Table Head

CP/70 Coral Bay

CP/71 Coral Bay

CP/72 Table Head

CP/73 Table Head

CP/74 Berkeley Bay

CP/75 Coral Bay

CP/76 Coral Bay

CP/77 Orontes Reef

CP/78 Orontes Reef

CP/80 Orontes Reef

CP/81 Kennedy Bay

CP/82 Coral Bay

CP/83 Coral Bay

CPV8 W\side of Barrow Bay

CP/85 Orontes Reef

CP/86 Table Head

CP/87 Victoria Settlement

CP/88 Orontes Reef

CP/90 Orontes Reef

CP/91 Orontes Reef

CP/92 Orontes Reef

CP/93 Coral Bay

CP/95 Coral Bay

CP/97 Coral Bay

CP/98 Coral Bay

CP/99 Orontes Reef

sia and northern Australia (Bruce 1980: col.
fig-)-

Urocaridella antonbruunii (Bruce)

Periclimenes antonbruunii Bruce, 1967: 45-
53, figs 19-22.

Leandrites cyrtorhynchus Fujino and Miyake
1969a: 143-149, figs 1-3; - Bruce 1983a: 42.

Urocaridella antonbruunii- Chace and Bruce
1993:43.

Material examined. 1 spm., stn CP/21, Coral
Bay, 1-3 m, in cave, 18 October 1981, coll. P.N.
Alderslade, H.K. Larson, J.R. Hanley, NTM
Cr.001311.

Remarks. This taxon was first reported as
Periclimenes antonbruunii on the basis of a
single small specimen from the Comoro Islands.

Date Habitat Depth

14m

4- 5m
LW
5m
LW
4m
<lm
<6m
4m
4m
3-4m
0.5m
2-3m

1- 4m

2- 5m
2-5m
3m
6-8m
3m
ca.5m
6 m
10m
19m
10m
2.5m

5- 6m
surface
LWS
10-15m
8m
LW

8-10m

12m

15-22m

6- 8m
5-6m
?

1 l-12m

10°59.0'S
11°14.8’S
11°14.8’S
11°14.7'S
11°14.2'S
11°14.4’S
11°13.9'S
11°11.0'S

ii°n.o’s

I rio.s's

11°12,0'S

lrn.o's

II “10.4'S
11°14.4'S

iru.2's

11°10.4’S
11°14.6’S
11°14.6’S
11°14.8’S

n°ii.rs

I l°11.2'S
11°04.5'S

II °04.5'S
11°04.5’S
11°12.8'S
n°it.rs
lrii.rs

11°21.5'S
ir04.5’S
11°I3.5’S
11°22.2'S
H°04.5'S
11“04.5'S
11°04.5'S
11°04.5'S
ll.l’S
11.0’S
11°11.2'S
11 °11,2’S
11°04.5'S

11 '

ll‘

132°48.7'E

132°10.8'E

132°11.2’E

132°10.2'E

I32°12.2'E

132°10.8’E

132°I2.3’E

132°03.4'E

132°03.8'E

132°03.0’E

I32°02.0'E

132°02.8'E

132°10.8'E

132°02.8’E

132°02.8'E

132°10.5'E

132 ,, 10.5’E

132°11.4'E

132°03.4'E

132°02.8'E

132 C 04.8'E

132°04.8’E

132°04.8'E

132°05.6'E

I32°03.4'S

132°13.0'E

132°04.75'E

I32°l 1.5'E

132°09.0'E

132°04.8'E

132°04.8’E

132°04.8'E

132°04.0'E

132°03.4’E

132°02.8’E

132°02.8'E

132°04.8’E

21.10.82

11.5.83

12.5.83

13.5.83

15.5.83

16.5.83

17.5.83

18.5.83

19.5.83
12.9.85

12.9.85

13.9.85

14.9.85

15.9.85

16.9.85

17.9.85

18.9.85
18.9.85

18.9.85

19.9.85

7.8.86

8.8.86

9.8.86

10 . 8.86
10.8.86
10.8.86
11.8.86
12.8.86
12.8.86
13.8.86
13.8.86

coral reef
coral & sand
basalt reef flat pond
rocky reef

sea grass bank in creek
coral reef
saltwater creek
coral reef edge
sheltered coral reef
coral reef
coral reef
sand beach
algal bank
coral reef
coral reef
coral reef

coral bommie
weedy reef

coral reef with fish pond
coral reef
off-shore bank
off-shore bank

seaward side of reef
coral reef
night-light
mud flat

rock bommie

coral reef
coral reef
coral reef
fine mud

coral reef, off fish pond
coral reef

rocky reef

103

A.J. Bruce and K.E. Coombes

104

Fig. 1. Map of the Cobourg Peninsula region, Arnhem Land, indicating collection localities referred to in text. l.TrepangBay. 2, Port Essington. 3, Port Bremer. 4, Raffles Bay.

The palaemonoid shrimp fauna of the Cobourg Peninsula

The specimen closely resembled larger adult
specimens subsequently collected, and was later
considered to be a juvenile, probably a first post-
larval stage, as rudimentary exopods werepresent
on the first and second pereiopods. The ambula¬
tory dactyls were also minutely biunguiculate, as
also occurs in the juveniles of some other
palaemonine shrimps, such as the dionyx stage
of Macrobrachium (Holthuis 1950). The third
abdominal segment was also more strongly
posterodorsally produced than in larger speci¬
mens. There have been no further reports of P.
antonbruunii under that name. The larger speci¬
mens were referred to Leandrites cyrtorhynchus
Fujino and Miyake (1969a), a species now known
to be of widespread distribution. The species is
now also well known as a fish cleaner. Both taxa
are characterised in life by a colourpattem of red
and white spots (Bruce 1980: col. fig.), and
enough immature specimens have been col¬
lected to present a continuous series, so that they
clearly represent a single species. Chace and
Bruce (1993) re-establish the genus Urocaridella
Borradaile, and removed this species from
Leandrites, placing it in Borradaile’s genus
Urocaridella, on account of the similarities of
the carapace and rostrum.

Australian distribution. Previously recorded
in the Northern Territory from Darwin Harbour
(Bruce 1983a). Also recorded from Heron Is¬
land, within the Great Barrier Reef (Bruce 1980).

Further distribution. Type locality: Pamanzi
Island, Mayotte, Comoro Islands. Also reported
from East Africa to Japan and New Caledonia.

Urocaridella sp. ?

Leandrites sp. Bruce, 1993: 36 (col. fig.).

Material examined. 4 spms, stn CP/20,
Walford Point,LWS, 170ctober 1981, coll. A. J.
Bruce etal., NTMCr.001338.1 spm., same data
as previous, NTM Cr.001339.9 spms, same data
as previous, Cr.001951. 2 spms, stn CP/21,
Coral Bay, in cave, 1-3 m, 18 October 1981, coll.
P.N. Alderslade et al., NTM Cr.001336. 10
spms, same data as previous, in cave, NTM
Cr.001337. 2o', stn CP/23, Walford Point, 5 m,
19 October 1981, NTM Cr.005242.

Remarks. This taxon is distinguished from
Urocaridella antonbruunii by its characteristic
colour pattern, which consists of conspicuous
yellow and black spots (Bruce 1993a). The pat¬
tern is consistent in all specimens, with minimal
variation, although less well developed in small
specimens. There are no intermediates with U.

Fig. 2. Map of PortEssington, Cobourg Peninsula, indicating
localities referred to in text.

antonbruunii. The shrimps look and behave like
a distinct species, but no morphological differ¬
ences have been detected so far that can be used
to separate the two taxa once the colour pattern
has been lost after preservation. The taxon is
therefore not provided with a specific name.

This species, and U. antonbruunii probably
represent a pair of closely related sibling spe¬
cies. A similar situation occurs in the taxa
represented by Coralliocaris venusta Kemp, in
which two distinctive colour patterns have been

designated aand p (Bruce 1979b) (the colouration

of the type material was not recorded, so that it
is not possible to associate either of these colour
patterns with the name designated by Kemp).
Specimens, in this case also, always segregate by
colour pattern. Other pairs of sibling species,
such as Periclimenes soror Nobili and P. bicolor
Edmondson, present similar taxonomic prob¬
lems and are reviewed in Bruce (1979b).
Harpiliopsis depressa (Stimpson) and H.
spinigera (Ortmann), long synonymised, were
first confirmed as distinct through the differ¬
ences in colour pattern in live specimens, and

105

A.J. Bruce and K.E. Coombes

morphological features were subsequently es¬
tablished that now enable the identities of pre¬
served material to be established. If live-colour
patterns are consistently noted, then the mor¬
phological separation of these closely related
taxa may become practicable.

Australian distribution. In addition to the
Port Essington specimens, this taxon has also
been found to occur in Darwin Harbour.

Subfamily Pontoniinae Kingsley, 1878

Anchistus australis Bruce

Anchistus australis Bruce, 1977: 56-62, figs
7-9; 1983a: 43-44.

Material examined. 4 spms, stn CP/5, Coral
Bay, 2-3 m, 23 June 1981, coll. J.N.A. Hooper,
A.J. Bruce, P. Horner, in the bivalve mollusc
Tridacna sp., NTM Cr.000091.

Australian distribution. Type locality: Capre
Cay, Swain Reefs, Queensland. Northern Terri¬
tory: Coral Bay, Port Essington (Bruce 1983a).
Queensland: Michaelmas Reef.

Further distribution. Also known from In¬
donesia; New Caledonia; Fijian Islands and
Marshall Islands.

Anchistus custos (Forsskal)

Cancer custos Forsskal, 1775: 94.

Harpilius inermis - Miers 1884: 291, pi. 32
fig. B.

Anchistus inermis - Borradaile 1898: 387; -
Kemp 1922: 249-252, fig. 81.

Anchistus custos - Holthuis 1952: 105-109,
figs 43-44; - Bruce 1983a: 44; 1988b:227.

Material examined. 2 spms, stn CP/5, Coral
Bay, 2-3 m, 23 June 1981, coll. P. Homer,
J.N.A. Hooper, A.J. Bruce, in the bivalve mol¬
lusc Pinna sp., NTM Cr.000095. 1 ovig. 9 . Id",
stn CP/ 8 , Coral Bay, 2-3 m, 24 June 1981, coll.
A.J. Bruce, J.N.A. Hooper, P. Homer, in Pinna
sp., NTM Cr.008549. 1 ovig. 9 , lei", 2 juv., stn
CP/17, Wanaray Point, Trepang Bay, LWS, 15
October 1981, coll. A.J. Bruce, P. Homer, NTM.
Cr.008728. 1 <f, 1 ovig. 9 , stn CP/18, Midjari
Point, Trepang Bay, LWS, 16 October 1981,
coll. A.J. Bruce, NTM Cr.007591. 3d", 39 , stn
CP/20, Walford Point, Coral Bay, LWS, 17
October 1981, coll. A.J. Bruce et al., in Pinna
sp., NTM Cr.000133. 2 spms, stn CP/21, Coral
Bay, 1-3 m, 18 October 1981, coll. P.N.
Alderslade et al., in Pinna sp., NTM.
Cr.001878. 2 spms, stn CP/21, Sandy Island

No. 2, 10 m, 21 October 1981, coll. J.N.A.
Hooper et al., in Pinna sp., NTM Cr.003202.
Id", I 9 , stn CP/28, Sandy Island No. 2, 6-7 m,
22 October 1981, coll. J. R. Hanley, in Pinna
sp., NTM Cr.000163. lo", I 9 , stn CP/33, Port
Bremer ,6 m, 1 May 1982, coll. P.N. Alderslade,
in Pinna bicolor, NTM Cr.000298. 2 spms, stn
CP/36, Sandy Island No. 2, 13 m, 2 April 1982,
coll. J.N.A. Hooper, in. Pinna sp., NTM
Cr.000412. 2 spms, stn CP/37, Table Head, 1-3
m, 3 May 1982, coll. H.K. Larson, in Pinna sp.,
NTM Cr.002960. 6 spms. stn CP/38, Table
Head, 2-4 m, 4 May 1982, coll. H.K. Larson et
al., in Pinna sp., NTM Cr.001962. Id", I 9 , stn
CP/44, Table Head, 4-5 m, 11 May 1983, coll.
N.L. Bruce, in large Pinna sp., NTM Cr.009408.
2 spms, stn CP/60, Coral Bay, < 6 m, 16 May
1983, coll. H.K. Larson, in Pinna sp.,NTM Cr.
001740. 2 spms, same data as previous, NTM
Cr.010256. lo", 1 ovig. 9 , stn CP/ 68 , Coral Bay,
2-3 m, 19 May 1983 coll. J.R. Hanley, N.L.
Bruce, in Pinna sp., NTM Cr.009283. 2 spms,
same data as previous, NTM Cr.010241.2 spms,
stn CP/69, Table Head, 1-4 m, 11 September
1985, coll. A.J. Bruce, in Pinna sp., NTM
Cr.003219.2 spms, stn CP/71, Coral Bay, LWS,
13 September 1985, coll. A.J. Bruce, in Pinna
sp., NTM Cr.003226. 1 juv., stn CP/73, Table
Head, 6-8 m, 14 September 1985, coll. C. Hood
et al., in Pinna sp., NTM Cr.007559. 3 <f, 3
ovig. 9 , same data as previous, NTM Cr.007560.
1 spm., stn CP/82, Coral Bay, 5-6 m, coll. R.
Williams. P.N. Alderslade, in Pinna sp., NTM
Cr.007575. Id", 1 ovig. 9 , sin CP/ 86 , Table Head,
8 m, 7 August 1986, C. Johnson et al., in Pinna
deltoides. NTM Cr.004097. 4 spms, stn CP/93,
Coral Bay, 6-8 m, 11 August, 1986, coll. S.
Slack-Smith, in Pinna bicolor, NTM
Cr.004131. 4 spms, same data as previous., in
Pinna deltoides, NTM Cr.004144. 2 spms, same
data as previous, NTM Cr.004147.

Australian distribution. Northern Territory:
Port Essington (Bruce 1983a), Darwin Harbour,
East Point (Bruce 1988b). Queensland: Port
Denison, Lizard Island, Magnetic Island, Swain
Reefs, Bowen, Port Molle. South Australia: St.
Vincent Gulf. Western Australia: Shark Bay,
Monte Bello Islands.

Further distribution. Type locality: Loheia,
Yemen. Common and widespread throughout
most of the Indo-West Pacific region from the
Red Sea, East Africa to Mozambique, to Japan,
east to the Philippines; Solomon Islands and
Fijian Islands.

106

The palaemonoid shrimp fauna of the Cobourg Peninsula

Chernocaris placunae Johnson

Chernocaris placunae Johnson, 1967: 500-
511, figs 1-12;-Bruce 1983a: 44; 1990b: 10.

Material examined. 6 o”, 6 ovig. 9 , stn HL81-
22, Arafura Sea, 10°58’S 132°10’E, 27 m, 19
October 1981, trawl, F.V. Anson, coll. A.J.
Bruce, NTM Cr.000102.

Remarks. The specimens have been previ¬
ously noted by Bruce (1983a), representing only
the second occurrence of this species. Unfortu¬
nately the locality of capture was incorrectly
given and the correct position is as given above.
As with the type material, the specimens were
found in association with the bivalve mollusc
Placuna placenta L.

Australian distribution. Arafura Sea,
10°58’S 132°10’E.

Further distribution. Type locality: Telok
Paku, Singapore. No further records.

Conchodytes monodactylus Holthuis

Conchodytes monodactylus Holthuis, 1952:
200-204, figs 96-98; - Bruce 1983a: 44.

Material examined. 1 ovig. 9 , lo", stn CP/29,
Sandy Island No. 2, 8-9 m, 22 October 1981,
coll. P. Horner, P.N. Alderslade, in bivalve
mollusc Pinna sp., NTM Cr.000092. Id”, I 9 , stn
CP/30, Black Point, 10-12 m, 29 April 1982,
coll. H.K. Larson, P. Homer, in Pinna sp. cf.
bicolor, NTM Cr.000296. 2 spms, stn CP/91,.
Orontes Reef, 10 August 1986, coll. S. Slack-
Smith, NTM Cr.004124.

Australian distribution. Northern Territory:
Sandy Island No. 2 (Bruce 1983a). Queensland:
Magnetic Island.

Further distribution. Type localities: Takao,
Taiwan; Lesser Sunda Islands, Indonesia. Also
known from Singapore and Hong Kong.

Coralliocaris graminea (Dana)

Oedipus gramineus Dana, 1852:25; 1855, pi.
37 fig. 3.

Coralliocaris graminea - Stimpson 1860:38;
- Bruce 1983a: 45.

Material examined. 1 spm., stn CP/ 8 , Coral
Bay, 2-3 m, 24 June 1981, parasitised by
Hemiarthrinae bopyrid (NTM Cr.000049), coll.
A.J. Bruce et al., NTM Cr.010751.1 spm., same
data as previous, bopyridized by Hemiarthrinae
(NTM Cr.000050), coll. A.J. Bruce et al., NTM
Cr.010752. la”, stn CP/17, Wanaray Point Reef,
Trepang Bay, LWS, 15 October 1981, coll. A.J.

Bruce, P. Homer, NTMCr.008731. Id”, 2 ovig. 9 ,
stn CP/20, Walford Point, Coral Bay, LWS, 17
October 1981, coll. A.J. Bruce et al., NTM
Cr.000118.1 spm., stn CP/21, Coral Bay, 1-3 m,
18 October 1981, coll. P.N. Alderslade et al.,
NTM Cr.001879. 3 spms (I 9 ,1 juv.), stn NY/9,
Oxley Island, LWS, 20 October 1982, on coral
Acroporasp,co\\. A.J. Bruce, NTM Cr.003165.
2 spms, stn CP/ 88 , Orontes Reef, 8-10 m, 9
August 1986, coll. C. Johnson, R.C. Willan, on
Acropora sp., NTM Cr.004103. 1 spm., stn CP/
90, Orontes Reef, 12 m, 10 August 1986, coll.
R.C. Willan, P. Davie, on Acropora sp., NTM
Cr.004115.

Australian distribution. Northern Territory:
Port Essington (Bruce, 1983a). Queensland:
Heron Island, Myora, Palm Island, Falcon Is¬
land, Willis Island, Bet Reef, Restoration Rock.

Further distribution. Type locality: Rewa,
Viti Levu, Fiji. Commonly and widely distrib¬
uted, from the Red Sea and East Africa to the
Bonin Islands, and Society Islands.

Coralliocaris viridis Bruce

Coralliocaris viridis Bruce, 1974: 22-224, fig.

1 .

Material examined. I 9 , 3 ovig. 9 , stn CP/5,
Coral Bay, 2-3 m, 23 June 1981, coll. A.J. Bruce,
P. Homer, J.N.A. Hooper on coral Acropora sp.,
NTM Cr.008562.1 ovig. 9 , stn CP/ 8 , Coral Bay,
2-3 m, 24 June 1981, coll. A.J. Bruce, P. Homer,
J.N.A. Hooper, on Acropora sp., NTM
Cr.008538. la”, same data as previous, NTM
Cr.008543. 1 ovig. 9 , same data as P revious ’
NTM Cr.008563. 5 spms (2 ovig. 9 ), stn CP/37,
Table Head, 1-3 m, 3 May 1982, on Acropora
sp., coll. A .J. Bruce, NTM Cr.000303.7 spms (2
ovig. 9 , 2 juv.), stn CP/40, Orontes Reef, 3 m, 5
May 1982, coll. H.K. Larson, J.N.A. Hooper,
NTM Cr.001236.3 spms, same data as previous,
coll. A.J. Bruce, NTM Cr.008776. Id", 4 ovig. 9 ,
1 juv., stn NY/10, Oxley Island, 14 m, 21
October 1982, coll. A.J. Brace et al., NTM
Cr.007736.3 spms (1 ovig. 9 ), stn CP/77, Orontes
Bay, 10 m, 16 September 1985, coll. A.J. Brace,
on Acropora valida, NTM Cr.007558. 1 ovig. 9 ,
same data as previous, NTM Cr.007633.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island, One Tree Island.

Further distribution. Type locality: Mom¬
basa, Kenya. Also known from Sri Lanka;
Maidive Islands; Mozambique Channel; Ryukyu

107

A.J. Bruce and K.E. Coombes

Islands; Misool Island, Indonesia and Madang,
Papua New Guinea.

Dasella ansoni Bruce

Dasella ansoni Bruce, 1983b: 22-28, figs 1-5;
1990b: 10.

Material examined. 1 ovig .9 holotype, lcf
allotype, 1 bopyridized 9 , stn HL81 -22, Arafura
Sea, 10°58’S 132° 10’E, 27 m,FV Anson, trawl,
19 October 1981,coll. A.J. Bruce, in an ascidian,
NTM Cr.000104.

Remarks. This species is still known only
from the type specimens, which were collected
during the Cobourg Peninsula survey, from a
locality to the NW of Orontes Reef. The position
of the type locality was incorrectly given in the
original description and is as given above. The
specimens were found in association with the
tunicate Phallusiadepressiuscula (Heller). There
have been no further collections of this species.

Australian distribution. Type locality:
Arafura Sea, 10°58'S 132°10’E.'

Further distribution. Not yet reported from
extra-Australian waters.

Hamodactylus boschmai Holthuis

Hamodactylus boschmai Holthuis, 1952:209-
212, figs 102-104; - Bruce 1988b: 227.

Material examined. 1 ovig. 9 , stn CP/28,
Sandy Island No. 2, 6-7 m, 22 October 1981,
coll. J.R. Hanley, NTM Cr.000162.

Australian distribution. Northern Territory:
East Point, Darwin (Bruce 1983a, 1988b).
Queensland: Heron Island.

Further distribution. Type localities: Temate,
Aru Islands, Indonesia. Also known from Zan¬
zibar; Kenya; Madagascar; Singapore; Hong
Kong and New Caledonia.

Hamodactylus noumeae Bruce

Hamodactylus noumeae Bruce, 1970b: 539-
541, fig. 2.

Material examined. 8 spms, stn CP/20,
Walford Point, Coral Bay, LWS, 17 October
1981, coll. A.J. Bruce, on gorgonian Rumphella
aggregata, NTM Cr.001942. Id", stn NY/2,
New Year Island, 10 m, 14 October 1982, coll.
A.J. Bruce, NTM Cr.009298.5 spms, stn NY/5,
McCluer Island, 1 m. 160ctober 1982, coll. A.J.
Bruce, P. Horner, NTM Cr.009328. 4 spms,
same data as previous, NTM Cr.009329. lcf, 1

ovig. 9 , stn NY/ 8 , Oxley Island, LWS, 19 Octo¬
ber 1982. coll. A.J. Bruce, NTM Cr.005630. 1
spm., same data as previous, on unidentified
gorgonian, NTM Cr.006360.5 spms, stn CP/62,
Walford Point, Coral Bay, 4 m, 17 May 1983,
coll. N.L. Bruce, J.R. Hanley, on gorgonian
Junceella, NTM Cr.010523.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island, Lizard Island.

Further distribution. Type locality: Noumea,
New Caledonia. Also known from Zanzibar,
Tanganyika; Kenya; Temate, Indonesia and
Okinawa, Ryukyu Islands.

Hamopontonia corallicola Bruce

Hamopontonia corallicola Bruce, 1970a: 41-
48, figs 1-4; 1987a: 166; 1988b: 227.

Material examined. 1 spm., stn CP/21, Coral
Bay, 1-3m, 180ctober 1981,coll. P.N.Alders-
ladec/a/.,NTMCr.001882. 1 spm., stn CP/37,
Table Head, 1-3 m, 3 May 1982, coll. A.J. Bruce,
NTM Cr.000302. 2 ovig. 9 , same data as previ¬
ous. on coral Pocillopora sp., coll. H.K. Larson,
NTM Cr.002939. lcf, stn CP/44, Table Head, 4-
5 m, 11 May 1983, coll. N.L. Bruce, NTM
Cr.009348.1 juv., stn CP/60, Coral Bay, < 6 m,
16 May 1983, coll. N.L. Bruce, NTM Cr.009281.
1 ovig. 9 , stn CP/64, Walford Point, Coral Bay,
3-4 m, 18 May 1983, coll. N.L. Bruce, on coral
Heliofungia actiniformis, NTM Cr.000448. 1
spm., stn CP/71, Coral Bay, 2-5 m, 13 Septem¬
ber 1985, coll. A.J. Bruce, on Heliofungia ac¬
tiniformis, NTM Cr.003225. lcf, stn CP/75, Coral
Bay, ca. 5 m, 15 September 1985, coll. J.E.N.
Veron, R. Williams, NTM Cr.007567.8 spms (1
ovig. 9 ), same data as previous, on coral Euphyl-
lia sp., NTM Cr.007637.1 spm., stnCP/82, Coral
Bay, 5-6 m, 18 September 1985, coll. R. Wil¬
liams, P.N. Alderslade, on Heliofungia actini¬
formis, NTM Cr.007580. 1 ovig. 9 , same data as
previous, NTM Cr.007574. lcf, 1 ovig. 9 , stn CP/
97, Coral Bay, 12 August 1986, coll. C. Johnson,
on Heliofungia actiniformis, NTM Cr.004167.

Australian distribution. Northern Territory:
Port Essington (Bruce 1987a); East Point, Dar¬
win (Bruce 1988b). Queensland: Heron Island,
Peloris Island.

Further distribution. Type locality: Kat O
Chau, Hong Kong. Also known from SE Honshu;
Kushimoto, Japan; Ryukyu Islands, Japan and
Banda Nura Island, Indonesia; (also Kei Is¬
lands?).

108

The palaemonoid shrimp fauna of the Cobourg Peninsula

Hamopontonia essingtoni Bruce

Hamopontonia essingtoni Bruce, 1987a: 158-
166, figs lc, 11-14, 15 d-g; 1990b: 11.

Material examined. 1 spm., stn CP/70, Coral
Bay, 2-5 m, 12 September 1985, coll. P.N.
Alderslade, C. Hood, on coral Stylophora sp.,
NTM Cr.007554.1 spm., same data as previous,
on coral Stylophora pistillata, NTM Cr.007621.
1 ovig .9 holotype, lo" allotype, stn CP/71, Coral
Bay, 2-5 m, 13 September 1985, coll. L. Vail, on
Stylophora pistillata, NTM Cr.004072.16 spms
(3 ovig. 9 ), same data as previous, coll. A.J.
Bruce, NTM Cr.007568. lo", same data as pre¬
vious, coll. A.J. Bruce, on Stylophora pistillata,
NTM Cr.007628. 12 spms (6 ovig. 9 ) paratypes,
stn CP/76, Coral Bay, 6 m, 15 September 1985,
coll. L. Vail, on Stylophora pistillata, NTM
Cr.004073 A and B. 1 spm., stn CP/82, Coral
Bay, 5-6 m, 18 September 1985, coll. P.N.
Alderslade, R. Williams, on Stylophora sp.,
NTM Cr.007608.

Australian distribution. Type locality: Coral
Bay, Port Essington (Bruce 1987a). No further
records.

Further distribution. Not known outside
Australian waters.

Harpiliopsis beaupresii (Audouin)

Palaemon beaupresii Audouin, 1825: 91.

Harpilius beaupresii - Heller 1861: 27.

Harpiliopsis beaupresi - Borradaile 1917:
324, 379, pi. 55, fig. 21; - Holthuis 1952; 181-
182.

Material examined, lo", 1 ovig. 9 , stn CP/16,
Trepang Bay, 2 m, 14 October 1981, coll. A.J.
Bruce et al., on coral Seriatopora sp., NTM
Cr.001901. I 9 , same data as previous, on coral
Stylophora sp., NTM Cr.008647. 1 spm., stn
NY/1, New Year Island, 10 m, 13 October 1982,
coll. P. Horner, on Seriatopora sp., NTM
Cr.003175.1 spm., same dataas previous, NTM
Cr.007744. lo", 1 ovig. 9 , same data as previous,
NTM Cr.007754. lo", 2 ovig. 9 , stn NY/2, New
Year Island, 10 m, 14 October 1982, coll. A.J.
Bruce, NTM Cr.009291. 2 ovig. 9 , 1 juv., stn
NY/3, New Year Island, 16m, MOctober 1982,
coll. P. Homer et al., NTM Cr.007775. 1 spm.,
stn NY/4, McCluer Island, 7 m, 16 October
1982, coll. A.J. Bruce, NTM Cr.009280. Id",
same data as previous, NTM Cr.009295. Id", 1
ovig. 9 , stn NY/5, McCluer Island, 1 m, 16
October 1982, coll. A.J. Bruce, P. Horner, NTM
Cr.009320. 1 ovig. 9 , same data as previous,

NTM Cr.009322. 7 spms (1 ovig. 9 ), stn NY/9,
Oxley Island, LWS, 20 October 1982, coll. A.J.
Bruce, on Seriatopora sp., NTM Cr.003168.
lcf, 1 ovig. 9 , same data as previous, NTM
Cr.009294. Id", 1 ovig. 9 , stn CP/71, Coral Bay,
2-5 m, 13 September 1985, coll. A.J. Bruce, on
Stylophora sp., NTM Cr.007561. Id", 1 ovig. 9 ,
same data as previous, NTM Cr.007565. lo", stn
CP/75, Coral Bay, ca. 5 m, 15 September 1985,
coll. J.E.N. Veron, R. Williams, on Stylophora
sp., NTM Cr.007638. 2d", 3 ovig. 9 , stn CP/77,
Orontes Bay, 10 m, 16 September 1985, coll. R.
Williams, on Stylophora pistillata, NTM
Cr.007555. 1 spm., same data as previous, on
Stylophora pistillata, NTM Cr.007632. Id", same
data as previous, on Seriatopora hystrix, NTM
Cr.007634. lo", 1 ovig. 9 , stn CP/82, Coral Bay,
5-6 m, 18 September 1985. coll. P.N. Alderslade,
R. Williams, 5-6 m, on Stylophora sp., NTM
Cr.007616. 1 spm., stn CP/95, Coral Bay, 12
August 1986, 5-6 m, coll. R. Williams et al.,
NTM Cr.004155.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island.

Further distribution. Type locality: Egyp¬
tian Red Sea. Abundant throughout the Indo -
West Pacific region: Red Sea; East Africa: In¬
dian Ocean islands; Indonesia; South China
Sea; Thailand; Japan; Philippines; Marshall
Islands; and Hawaiian Islands, also extending to
Easter Island.

Ischnopontonia lophos (Barnard)

Philarius lophos Barnard, 1962: 242-243,
f'g- 2 .

Ischnopontonia lophos - Bruce 1966: 585-
595, figs 1-5; - Bruce 1983a: 44; 1988b: 227-
228.

Material examined. Id", 1 ovig. 9 , stn NY/ 8 ,
Oxley Island, LWS, 19 October 1982, coll. J.
Robinson, A.J. Bruce, on coral Galaxea
fascicularis, NTM Cr.007760. 3 spms, stn CP/
75, Coral Bay, 5 m, 15 September 1985, coll.
J.E.N. Veron, R. Williams et al., on Galaxea
fascicularis, NTM Cr.007639.

Australian distribution. Northern Territory:
East Point, Darwin (Bruce 1983a, 1988b); Port
Essington. Queensland: Heron Island, One Tree
Island, Great Palm Island, Orpheus Island,
Fan tome Island.

Further distribution. Type locality: Inhaca
Island, Mozambique. Also known from Zanzi¬
bar; Kenya; Tanganyika; Comoro Islands; Mada-

109

A.J. Bruce and K.E. Coombes

gascar; Seychelle Islands; Malaya; Singapore;
Caroline Islands; Fijian Islands and Ryukyu
Islands.

Onycocaris quadratophthalma (Balss)

Pontonia quadratophthalma Balss, 1921: 15,
fig. 7.

Onycocaris quadratophthalma - Holthuis
1952: 150-151;-Bruce 1993b: 330-335,figs 1-3.

Material examined. 1 ovig. 9 , stn CP/37,
Table Head, 1-3 m, 3 May 1982, coll. A.J. Bruce,
in unidentified sponge, NTM Cr.000301.

Remarks. Onycocaris quadratophthalma ap¬
pears to represent a complex of related species
and most of the non-Australian records require
detailed re-examination. Outside Australia,
known with certainty only from Hong Kong.

Australian distribution. Type locality: Cape
Jaubert, Western Australia. Northern Territory:
not previously recorded.

Further distribution. Reported only from
Eniwetak Atoll, Marshall Islands; Hong Kong;
Wake Island; Pearl and Hermes Reefs; Oahu,
Hawaiian Islands, and possibly also Ryukyu
Islands.

Palaemonella pottsi (Borradaile)

Periclimenes (Falciger) pottsi Borradaile,
1915: 212.

Palaemonella pottsi - Kemp 1922: 126-127;
-Bruce 1988b: 228.

Material examined, lei", 1 ovig. 9 , stn CP/26,
Sandy Island No 2, 7 m, 20 October 1981, coll.
P.N. Alderslade, J.N. A. Hooper, on unidentified
crinoid, NTM Cr.010108.1 spm., stn NY/1, New
Year Island, 10 m, 13 October 1982, coll. P.
Homer, on unidentified crinoid, NTM Cr.007751.
lei", stn NY/4, McCluer Island, 7 m, 16 October
1982, coll. AJ. Bruce, NTM Cr.009288. 1 spm.,
stn CP/62, Coral Bay, 4 m, 17 May 1983, coll.
N.L. Bruce, J.R. Hanley, on crinoid Comanthina
schlegelii , NTM Cr.000446. lei", 1 ovig. 9 , stn
CP/76, Coral Bay, 6 m, 15 September 1985, coll.
L. Vail, J.E.N. Veron, R. Williams, on crinoid
Stephanometra oxyacatitha , NTM Cr.007625.
I 9 , stn CP/80, Orontes Bay, 10 m, 17 September
1985, coll. J.E.N. Veron, R. Williams.on crinoid
Comanthus briareus, NTM Cr.007627. 2 spms,
stn CP/85, Orontes Reef, 10-15m, 19September
1985, coll. C Hood, L. Vail, R. Williams, on
unidentified crinoid, NTM Cr.007614.

Remarks. Palaemonella pottsi is known to
associate with a wide variety of crinoid hosts, but

has not been previously reported in association
with the genus Stephanometra.

Australian distribution. Type locality:
Mabuaig, Torres Strait, Queensland. Northern
Territory: East Point, Darwin (Bruce 1988b).
Queensland: One Tree Island, Heron Island,
Capricorn Islands.

Further distribution. Zanzibar; Singapore;
Indonesia; Japan; New Caledonia; Philippines;
and Marshall Islands.

Palaemonella rotumana (Borradaile)

Periclimenes rotumanus Borradaile, 1898:
383.

Palaemonella vestigialis Kemp 1922: 123-
126, figs 1-2, pi. 3, fig. 2.

Palaemonella rotumana - Bruce 1970c: 276-
279: pi. 1 e-f; 1983a: 42; 1988b: 228.

Material examined, la", stn CP/2, Kennedy
Bay, 2 m, 22 June 1981 , coll. A.J. Bmce, J.N.A.
Hooper, P. Homer, on unidentified sponge, NTM
Cr.008542. 10", same data as previous, on coral
Acropora sp., NTM Cr.008546.2 spms (1 ovig. 9 ),
same data as previous, on Acropora sp., NTM
Cr.008557. 29 , stn CP/5, Coral Bay, 2-3 m, 23
June 1981, coll. A.J. Bruce, J.N.A. Hooper, P.
Horner, on Acropora sp., NTM Cr.008532. I 9 ,
same data as previous, NTM Cr.008533.1 ovig. 9 ,
same data as previous, NTM Cr.008534. 1 spm.,
stn CP/ 8 , Coral Bay, 2-3 m, 24 June 1981, coll.
A.J. Bruce, J.N.A. Hooper, P. Horner, on
Acropora sp., NTM, Cr.008545.1 spm., stn CP/
10, Black Point, 1-2 m, 18 July 1981, coll. A.J.
Bruce, J.N.A. Hooper, on brown algae Padina
sp., NTM Cr.007653. 19 , same data as previous,
on coral Turbinaria sp., NTM Cr.007656. lo",
same data as previous, on Padina sp., NTM
Cr.007657. lo", same data as previous, in rub¬
ble, NTM Cr.007658. lo", 1 ovig. 9 , same data as
previous, on Turbinaria sp., NTM Cr.007661.2
spms, stn CP/14, Burford Island, LWS, 13 Octo¬
ber 1981, coll. A.J. Bmce etal.. NTM Cr.003192.
Id", 1 ovig. 9 , stn CP/16, Trepang Bay, 2 m, 14
October 1981, coll. A.J. Bmce et al., on coral
Stylophora sp., NTM Cr.(>08646.1 ovig. 9 ,2 juv.,
stn CP/17, Wanaray Point, Trepang Bay, LWS,
15 October 1981, coll. A.J. Bmce, P. Homer,
NTM Cr.008711 . l 9 juv„ stn CP/30, Black Point,
10-12 m, 29 April 1982, coll. P. Homer, H.K.
Larson, NTM Cr.009326. lo", stn CP/34, Dan¬
ger Point, Port Bremer, 2 m, 1 May 1982, coll.
J.R. Hanley, R. Williams, NTM Cr.009332. lo",
I 9 , stn CP/36, Sandy Island No 2, 13 m, 2 May
1982, coll . J.N. A. Hooper etal. , NTM Cr.009333.

110

The palaemonoid shrimp fauna of the Cobourg Peninsula

2 spms, stn CP/38, Table Head, 2-4 m, 4 May

1982, coll. H.K. Larson etal., NTM Cr.001966.
Id", stn NY/2, New Year Island, 10 m, 14 Octo¬
ber 1982, coll. AJ. Bruce, NTM Cr.009315. la",
1 ovig. 9 , stn NY/3, New Year Island, 16 m, 14
October 1982, coll. P. Horner et al., NTM
Cr.007776.2 ovig. 9 , stn NY/ 8 , Oxley Island, 19
October 1982, coll. J. Robinson, A.J. Bruce, NTM
Cr.007758.1 spm., same data as previous, NTM
Cr.007767. 1 ovig. 9 , stn CP/46, off Table Head,
5 m, 12 May 1983, coll. N.L. Bruce, NTM
Cr.009356.1 spm., stn CP/51, Table Head, 4 m,
13 May 1983, coll. N.L. Bruce, on Acroporasp.,
NTM Cr.010481. 1 spm., stn CP/62, Coral Bay,
4 m, 17 May 1983, coll. J.N.A. Hooper, on uni¬
dentified sponge, NTM Cr.00044 L 3 spms (1
ovig. 9 ), stn CP/64, Coral Bay, 3-4 m, 18 May

1983, coll. N.L. Bruce, on Acropora sp., NTM
Cr.009279.1 ovig. 9 , same data as previous, NTM
Cr.010524.2o\ I 9 , stn CP/65, Middle Bay, Coral
Bay, 0.5 m, 18 May 1983, coll. A.J. Bruce, NTM
Cr.009276. 1 spm., stn CP/75, Coral Bay, ca. 5
m, 15 September 1985, coll. J.E.N. Veron, R.
Williams, on Acropora sp., NTM Cr.007605. 1
spm., same data as previous, on coral Galaxea
fascicularis, NTM Cr.007636. lei", stn CP/82,
Coral Bay,5-6 m, 18 September 1985, coll. P.N.
Alderslade, R. Williams, on Acropora sp., NTM
Cr.007578. lo", same data as previous, on
Acropora digitifcra , NTM Cr.007611. 1 spm.,
same data as previous, on Stylophora sp., NTM
Cr.007617. lo", 1 ovig. 9 , sm CPV/ 8 , Barrow Bay,
LWS, 18 September 1985, coll. J.R. Hanley et
al., mangrove mudflats, NTM Cr.007648. 2
ovig. 9 , same data as previous, NTM Cr.007649.

Australian distribution. Northern Territory:
East Point, Weed Reef, Darwin Harbour (Bruce
1983a, 1988b). Queensland: Low Isles, Moreton
Bay, Heron Island, One Tree Island, North East
Cay, Herald Islands.

Further distribution. Type locality: Rotuma
Island. Common throughout most of the Indo-
West Pacific region, including Red Sea and East
Africa to Ryukyu and Hawaiian Islands, spread¬
ing via the Suez Canal into eastern Mediterra¬
nean Sea, but not reported from Eastern Pacific
region.

Palaemonella spinulata Yokoya

Palaemonella spinulata Yokoya, 1936: 135,
fig. 4.

Material examined. 1 spm., stn CP/30, Black
Point, 10-12 m, 29 April 1982, coll. P. Homer,
H.K. Larson, NTM Cr.000297.1 ovig. 9 , stn CP/

85, Orontes Reef, 10-15 m, 19 September 1985,
coll. C. Hood et al., NTM Cr.004056.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Moreton
Bay, Heron Island.

Further distribution. Type locality: Misaki,
Japan. Otherwise known only from Kenya, Tan¬
ganyika and La Reunion.

Periclimenaeus arabicus (Caiman)

Periclimenes (Periclimenaeus) arabicus
Caiman, 1939: 210-211, fig. 4.

Periclimenaeus arabicus - Holthuis 1952: 130.

Periclimenaeus ohshimae Miyake and Fujino
1967: 275-279, fig. 1.

Material examined. 2 ovig. 9 , sm CP/44, Table
Head, 4-5 m, 11 May 1983, coll. N.L. Bruce, in
sponge Callyspongia diffusa, NTM Cr.009409.

Australian distribution. Northern Territory,
not previously recorded. Queensland: Heron
Island, Capricorn Islands.

Further distribution. Type locality: Oman,
19°22.6’N 57°53.0’E. Alsoknown from Jibouti;
Kenya; Zanzibar; Tanganyika; Maidive Islands;
Hong Kong; Vietnam; Japan; New Caledonia
and Fijian Islands.

Periclimenaeus orontes Bruce

Periclimenaeus orontes Bruce, 1987a: 151-
158, figs lb, 6-10; 1990b: 11.

Material examined. I ovig. 9 , holotype, stn
CP/40, Orontes Reef, 3 m, 5 May 1982, coll. A.J.
Bruce, in sponge Jaspis stellifera, NTM
Cr.000272.

Australian distribution. Type locality:
Orontes Reef, Cobourg Peninsula, Northern Ter¬
ritory. No other records.

Further distribution. Not known outside
Australian waters.

Periclimenaeus serrula sp. nov.

(Figs 3-5)

Type material. 1 ovig. 9 , holotype, lei" allotype,
stn CP/99, Orontes Reef, 11°4.5’S 132°4.8’E, 11-
12 m, SCUBA, 13 August 1986, coll. A. Hoggett
et al., in tunicate Leptoclinoides incertus, NTM
Cr.004174 A (holotype) & B (allotype).

Diagnosis. Medium sized shrimp for the ge¬
nus, female of swollen form, male slender.

Rostrum (Fig. 3c) well developed, reaching to
about end of proximal segment of antennular
peduncle, slender, upturned, dorsal carina obso-

111

AJ. Bruce and K.E. Coombes

Fig. 3. Periclimenaeus serrula sp. nov., ovigerous female holotype, Orontes Reef. A, anterior carapace, antennae lateral; B
same dorsal; C, rostrum; D, antennule; E, antenna; F, eye, dorsal; G, third maxilliped; H, sixth abdominal segment, dorsal; I
telson; J, same posterior margin; K, uropod.

112

The palaemonoid shrimp fauna of the Cobourg Peninsula

lete, with four dorsal teeth, first stout, well in
advance of orbital margin, others slender, distal
third without teeth, ventral border convex, un¬
armed, non-setose. Carapace (Fig. 3a, b) smooth,
without supraorbital spines ortubercles; inferior
orbital angle broad, convex; antennal spine well
developed, slender, acute, anteroventral angle

of branchiostegite bluntly produced. Abdomen
with first segment without dorsal lobe, sixth
segment (Fig. 3h) about 1.5 times broader than
long, posterolateral angles small, blunt. Telson
(Fig. 3i) 1.8 times longer than anterior width,
lateral margins feebly convex, convergent, dor¬
sal spines small, at 0.3 and 0.77 of telson length,

Fig. 4. Periclimenaeus serrula sp. nov., ovigerous female holotype. A, first pereiopod; B, same, chela, C, major second
pereiopod; D, same, chela; E, minor second pereiopod; F, same, chela; G, third pereiopod; H, same, dactyl, propod, distal carpus.

113

A.J. Bruce and K.E. Coombes

Fig. 5. Periclimenaeus serrula sp. nov.,ovigerous female holoty pe. A, major second pereiopod, fingers; B, same, tips of fingers;
C.cuttingedgeofdactyl; D, minor second pereiopod, fingers, medial; E, same lateral; F, third pereiopod, dactyl. Male allotype;
G, major second pereiopod, fingers; H, minor second pereiopod, fingers; I, third pereiopod, dactyl.

114

The palaemonoid shrimp fauna of the Cobourg Peninsula

anterior spines about 0.07 of telson length,
posterior spines slightly smaller, posterior mar¬
gin (Fig. 3j) broadly convex, about 0.4 of ante¬
rior width, lateral spines slightly smaller than
posteriordorsal spines, intermediate spines about
0.1 of telson length, slender, acute, submedian
spines similar, shorter, setulose.

Proximal segment of antennular peduncle
(Fig. 3d) with distolateral angle feebly pro¬
duced, with small acute tooth; stylocerite short,
broad, distally acute. Antenna (Fig. 3e) with
basicerite lacking dorsal lobe; scaphocerite reach-
ing to about middle of distal segment of
antennular peduncle, 2.6 times longer than broad,
lamina broadly rounded distally, far exceeding
small acute distolateral tooth. Eyes (Fig. 3f) with
small oblique hemispherical cornea.

Third maxilliped (Fig. 3g) with ischiomerus
and basis fused, with small functional
arthrobranch.

First pereiopod (Fig. 4a) moderately slender,
exceeding antennular peduncle by distal third of
merus; chela (Fig. 4b) with palm subcylindrical,
2.4 times longer than deep, fingers about 0.8 of
palm length, slender, spatulate, with bidentate
tips, cutting edges entire, dactylus without dor¬
sal setal tuft, carpus about 1.4 times chela length,
subequal to merus.

Second pereiopods (Figs 4c, e) grossly un¬
equal, dissimilar. Major chela (Fig. 4d) 1.6
times carapace length, 1.75 times minor chela
length, palm swollen, 1.8 times longer than
deep, smooth; dactyl (Fig. 5a) about 0.3 of palm
length, compressed, with large molar process,
tip acute (Fig. 5b), cutting edge (Fig. 5c) laminar,
minutely serrate; fixed finger with large fossa
proximally, with blunt tooth dorsally, cutting
edge blunt, entire, tip acute; merus without
spines or tubercles ventrally. Minor chela (Fig.
4f) about 1.75 of major chela length, subequal to
carapace length, palm about 2.0 times longer
than deep, feebly tapering distally, glabrous,
dactyl (Fig. 5d) about 0.37 of palm length, 2.4
times longer than deep, dorsal margin broadly
convex, cutting edge sublinear, sharply carinate,
unarmed, tip blunt; fixed finger short, about
0.25 of palm length, slightly exceeding half
dactyl length, deeply cannulate, tip strongly
hooked, acute; merus ventrally smooth.

Ambulatory pereiopods robust. Third
pereiopod (Fig. 4g) with propod (Fig. 4h) stout,
about 3.0 times longer than deep, tapering
strongly distally, with two stout distoventral
spines; dactyl (Fig. 50 small, compressed, about

1.5 times longer than deep, 0.2 of propod length,
unguis distinct, about 0.3 of dorsal corpus length,
corpus simple, without accessory teeth.

Uropod (Fig. 3k) with exopod laterally entire,
feebly setose, with small acute distolateral tooth
with mobile spine medially.

Remarks. Ova small about 0.5mm length.
The male is generally similar to female, rostral
dentition 4/0, body form slender, major chela
(Fig. 5g) about 1.95 times carapace length,
minor chela (Figs 5d, e) 0.9 times carapace
length, 0.46 of major chela length, ambulatory
dactyls similar to female, simple.

Etymology. From serra (latin), a saw, di¬
minutive, with reference to the cutting edge of
the dactyl of the major second pereiopod.

Host. Leptoclinoides incertus Sluiter
(Urochordata) (det. P. Mather, 20 August 1986).

Measurements. Ovigerous female, CL 4.5
mm; male, CL 2.5 mm.

Systematic position. Periclimenaeus serrula
is most closely related toP. crassipes (Caiman,
1939) and P. hecate (Nobili, 1904). Both these
species have simple unarmed dactyls on the
ambulatory pereiopods, lacking accessory teeth
or denticulations. P. serrula may be distin¬
guished from both these species by the presence
of an entire cutting edge on the dactyl of the
minor second pereiopod, and a minutely
denticulate distal cutting edge on the dactyl of
the major second pereiopod. The only other
Indo-West Pacific species with simple dactyls
on the ambulatory pereiopods are P.
arthrodactylus Holthuis, in which very large
supraorbital teeth are present, and P. trispinosus
Bruce, which has several post-antennal spines.

Periclimenaeus solitus sp. nov.

(Figs 6-7)

Type material. Id" holotype, stn CP/40,
Orontes Reef, 11°03.6’S 132°05.0 E, 3 m, 5
May 1982, coll. A.J. Bruce, in sponge Jaspis
siellifera , NTM Cr.000277.

Diagnosis. Small sized shrimp for the genus,
of slender subcylindrical body form.

Rostrum (Fig. 6c) well developed, slender,
acute, reaching to about distal end of proximal
segment of antennular peduncle, horizontal,
lateral carinae obsolete, dorsal carina with three
acute teeth on distal half, ventral margin un¬
armed, sinuous.

Carapace (Figs 6a, b) without supraorbital
spines or tubercles, with well marked postorbital

115

A.J. Bruce and K.E. Coombes

Fig. 6. Periclimenaeus solitus sp. nov., holotype male, Orontes Reef. A, carapace, eye and antennae, lateral; B, anterior
carapace, eyes, antennal peduncles, dorsal; C, rostrum and orbital region, lateral; D, antennule; E, antenna; F, eye, right, dorsal;
G, third pereiopod, distal propod and dactyl; H, sixth abdominal segment dorsal; I, male second pleopod, endopod; J, telson;
K, uropod.

116

The palaemonoid shrimp fauna of the Cobourg Peninsula

shoulder; antennal spine slender, acute, inferior
orbital angle small, anterolateral angle of
branchiostegite not produced, bluntly obtuse. Ab¬
domen with first segment lacking anterior dor¬
sal lobe; sixth segment (Fig. 6h) about 1.6 times
broader than long, posterolateral angles small,

acute. Telson (Fig. 6j) about twice as long as
anterior width, lateral margins feebly convex,
convergent, dorsal spines about 0.17 of telson
length, at 0.18 and 0.58 of telson length, subequal,
anterior pair submarginal, posterior pair mar¬
ginal, posterior margin broadly convex, about

117

A.J. Bruce and K.E. Coombes

0.5 of anterior margin width, with minute acute
median point, lateral spines about 0.6 of dorsal
spine length, intermediate spines 0.25 of telson
length, slender, very acute, submedian spines
0.9 of intermediate spine length, much more slen¬
der, setulose.

Proximal segment of antennular peduncle
(Fig. 6d) with distolateral margin produced,
with small acute lateral tooth; stylocerite short,
broad, acute. Antenna (Fig. 6e) with basicerite
lacking dorsal lobe; scaphocerite exceeding in¬
termediate segment of antennular peduncle,
about 2.7 times longer than broad, with small
acute distolateral tooth distinctly exceeding la¬
mella; carpocerite short, reaching to about 0.6
of scaphocerite length. Eyes (Fig. 6f) well de¬
veloped, with large oblique hemispherical cor¬
nea.

First pereiopod (Fig. 7a) moderately slender,
distal merus reaching almost to end of antennular
peduncle; chela (Fig. 7b) with palm
subcylindrical, slightly swollen, 2.5 times longer
than deep, fingers about 0.6 of palm length,
broadly subspatulae, lateral cutting edges entire,
tips expanded, bidentate, dactyl without dorsal
setal tuft; carpus about 1.2 times palm length,
subequal to merus.

Second pereiopods grossly unequal, dissimi¬
lar. Major chela (Fig. 7c) about 1.5 times cara¬
pace length, 1.5 times minor chela length, palm
subcylindrical, about 2.0 times longer than deep,
feebly tapered distally, compressed, dorsal sur¬
face acutely tuberculate, sparsely setose; dactyl
(Fig. 7d) about0.37 of palm length, compressed,
twice as long as deep, with broad acute tip, cut¬
ting edge with long, low molar process proxi-
mally, distal cutting edge entire; fixed finger
(Fig. 7d) with large fossa proximally, with large
acute tooth dorsally; ventral border of merus
strongly tuberculate (Fig. 7e), ischium feebly tu¬
berculate. Minor chela (Fig. 7f) about 0.7 of major
chela length, palm about 2.1 times longer than
broad, strongly compressed, acutely tuberculate,
with numerous long setae, especially ventrally;
dactyl (Fig. 7g) about 0.45 of palm length. 2.4
times longer than deep, with hooked bidentate
tip, dorsal margin strongly convex, cutting edge
very feebly convex, entire; fixed finger (Fig. 7g)
about 0.3 of palm length, distinctly exceeded by
dactyl, tip feebly acute, hooked, cutting edge
deeply cannulate; merus feebly tuberculate
ventrally.

Ambulatory pereiopods slender. Third
pereiopod (Fig. 7h) with propod (Fig. 7i) about

4.3 times longer than proximal depth, feebly
tapering distally, with four robust ventral spines,
two distoventral spines, medial spine longer
than lateral spine; feebly denticulate dorsally;
dactyl (Fig. 6g) about 0.28 of propod length,
slender, compressed, 3.0 times longer than deep,
unguis large, robust, curved, unarmed, about 0.8
of corpus length, corpus with strong, acute,
distally directed, distal accessory tooth, ventral
border with three minute denticles.

Male second pleopod (Fig. 6i) with appendix
masculina with short corpus, with two long
distal spines, 3.0 times corpus length, medial
spine setulose, lateral spine non-setulose.

Uropod (Fig. 6k) with exopod laterally entire,
sparsely setose, with small acute distal tooth
with mobile spine medially.

Host. Jaspis stellifera (Carter) (Jaspidae:
Porifera).

Etymology. From solitiis, latin, usual.

Measurement. Holotype male, CL 1.8mm.

Systematic position. Periclimenaeus solitus
is most closely related to Periclimenaeus
spongicola Holthuis, 1952. Features shared with
P. spongicola are: (i) third pereiopod dactyl
biunguiculate, not elongate, ca. 0.33 of propod
length, corpus with ventral denticulations; unguis
without ventral denticulations; (ii) exopod of
uropod laterally entire; (iii) dorsal margin of
first abdominal segment without anterior lobe;

(iv) dorsal telson spines not restricted to anterior
halfof telson; (v) dactyl of major second pereiopod
not subrectangular, (vii) first pereiopod fingers
greaterthan0.5ofpalm length; (viii) supraorbital
spines absent, (iv) scaphocerite much exceeding
carpocerite.

Periclimenaeus solitus may be distinguished
from P. spongicola by the following features:

(i) three instead of six dorsal rostral teeth;
rostrum straight, not up-curved; (ii) carpocerite
short, not exceeding scaphocerite; (iii)
scaphocerite with lamella not markedly exceed¬
ing distolateral spine; (iv) second pereiopod
with palms of chelae acutely tuberculate, not
smooth; merus ventrally tuberculate, not smooth;

(v) dactyl of minor second pereiopod with cut¬
ting edge entire, not dentate; (vi) third pereiopod
with propod slender, with strong ventral spines,
not swollen, with distoventral spines only; (vii)
third pereiopod dactyl slender, with anteroverted
distal accessory spine, not stout, with recurved
accessory spine; (viii) second pereiopods with
chelae tuberculate not smooth; merus ventrally
tuberculate, not smooth.

118

The palaemonoid shrimp fauna of the Cobourg Peninsula

Remarks. The holotype of Periclimenaeus
orontes was also obtained from a specimen of the
same host sponge, Jasp is stellifera, also from
Orontes Reef (Bruce 1987b). This species may
be readily distinguished by the presence of six
dorsal rostral teeth, with an anterior dorsal lobe

in the first abdominal tergite, and with the two
pairs of dorsal telson spines, both on the anterior
half of the telson. Jaspis stellifera is also host for
other pontoniine shrimps: Orthopontonia
ornatus (Bruce, 1970d) and Periclimenaeus
bidentatus Bruce, 1970d.

119

A.J. Bruce and K.E. Coombes

Periclimenaeus stylirostris Bruce
(Figs 8-9)

Periclimenaeus stylirostrisBruce, 1969:167-
168; 1972: 68-75, figs 2-6.

Periclimenaeus sp. Lowry and Springthorpe
1992: 129.

Material examined. Post-larva, stn CP/18,
Trepang Bay.LWS, 160ctober 1981,coll. A.J.
Bruce etal., NTM Cr.007667. Carapace length,
1.1 mm.

Description. Rostrum (Fig. 8b) slender, acute,
horizontal, reaching to about distal margin of
proximal segment of antennular peduncle, with
four small acute dorsal teeth, all well in advance
of posterior orbital margin, ventral margin un¬
armed. Carapace (Fig. 8a) without supraorbital
spines or tubercles, orbital notch large, antennal
spine well developed, anterolateral angle of
branchiostegite bluntly obtuse.

Telson (Fig. 8h) about 2.5 times longer than
proximal width, with two pairs of dorsal spines,
anterior pair subdorsal, about 0.12 of telson
length, at 0.15 of telson length, posterior pair
smaller, marginal, at 0.66 of telson length,
posterior margin feebly convex, about 0.45 of
anterior width, without median point, lateral
spines small, more slender than distal dorsal
spines, intermediate spines robust, 0.2 of telson
length, submedian spines slightly shorter,
setulose.

Antennular peduncle (Fig. 8c) with proximal
segment withdistolateral angle produced, acute,
feebly bifid on left, with ventromedial tooth,
stylocerite short, acute. Antenna (Fig. 8d) with
basicerite unarmed; scaphocerite exceeding
antennular peduncle, with acute distolateral
tooth, slightly exceeded by lamella, carpocerite
short, notreaching half scaphocerite length. Eye
large, cornea hemispherical, diameter about 0.3
of carapace length.

First pereiopod (Fig. 8e) short, stout; chela
subcylindrical, compressed, 1.5 times longer
than deep, fingers subequal to palm length,
robust, tapering, with small acute hooked tips,
cutting edges lateral, entire; carpus about 0.8 of
chela length; merus subequal to chela; ischium
about 0.7 of chela length.

Second pereiopods (Fig. 9a) subequal, simi¬
lar. Right chela (Fig. 9b) about 1.1 times cara¬
pace length, palm about 2.1 times longer than
proximal depth, tapering distally, moderately
compressed, generally smooth, with four rela¬
tively large very acute tubercles dorsally, one

ventrally; dactyl (Figs 9c, d) subcircular, about
1.25 times longer than deep, about 0.38 of palm
length, highly compressed, laminar, with strong
acute distal tooth, with convex entire sharp
cutting edge, without molar process; fixed fin¬
ger about 1.5 times longer than deep, tapering
distally to strongly acute bidentate tip, upper
tooth slightly long than lower, cutting surface
deeply cannulate, lateral margin strongly raised,
convex, lower margin concave; carpus normal,
distally excavate; merus about 0.5 of palm length,
unarmed; ischium 1.3 times meral length, un¬
armed. Left pereiopod (Fig. 9d) with fingers
similar, without molar process.

Ambulatory pereiopods robust. Third
pereiopod (Fig. 8f) with dactyl (Fig. 8g) about
0.3 of propod length, unguis well developed,
slender, unarmed, 3.0 times longer than basal
width, 0.65 of corpus length, corpus 1.6 times
longer than deep, compressed, feebly tapered
distally, with large slender, very acute
preterminal accessory tooth, ventral border oth¬
erwise unarmed; propod (Fig. 9e) about 0.45
times carapace length, 4,0 times longer than
deep, with one long distoventral spine, three
ventral spines. Fourth pereiopod similar, propod
(Fig. 9f) 0.85 of third propod length, with two
ventral spines. Fifth pereiopod with propod (Fig.
9g) slender, 6.0 times longer than deep, about
1.15 times longer than third propod, without
ventral spines; dactyl with accessory tooth fee¬
bly developed.

Uropod (Fig. 8i) without special features,
lateral margin of exopod feebly convex, entire,
with small acute distolateral tooth, with larger
mobile spine medially.

Remarks. The larval and post-larval stages of
the numerous Indo-West Pacific species of the
genus Periclimenaeus are at present all
undescribed. Gurney and Lebour (1941) de¬
scribed the larvae and post-larvae of some Ber¬
mudan species: Periclimenaeus (?)wilsoni (Hay),
Periclimenaeus sp. B. and Periclimenaeus
bermudensis (Armstrong); since then no further
descriptions have been published.

The genus Periclimenaeus at the moment
contains some 56 species, with 45 Indo-West
Pacific species, all presenting a wide range of
minormorphological variations at species level.
The morphology of the first post-larval stages
provides some indications of the plesiomorphic
state from which all these minor variations may
be derived. Even so, Periclimenaeus sp. B. of
Gurney and Lebour (1941) and the present speci-

120

The palaemonoid shrimp fauna of the Cobourg Peninsula

men show some close resemblances that are not
present in the post-larva of Periclimenaeus
wilsorti. In these two, the major second pereiopods
are subequal and similar, the fingers of the
chelae are similar in the twotaxa, with the fixed
finger deeply cannulate and the dactyl lacking a
distinct molar process (one of the diagnostic
features of the genus when adult). In the present

specimen, the dactyl is much deeper and the
fixed finger distally bidentate, in contrast to
Periclimenaeus sp. B. Periclimenaeus sp. B.
also differs from the present specimen in the
presence of rudimentary exopods on the first
four pereiopods. Periclimenaeus (?)wilsoni also
has these exopods (not shown in Gurney and
Lebour 1941: fig. 17k), but more significantly, it

Fig. 9. Periclimenaeus stylirostris, post-larva. A, second pereiopod, left; B, right, chela; C, same, fingers, D, left, distal fingers
of chela; E, third pereiopod, propod and dactyl; F, fourth pereiopod, same; G, fifth pereiopod, same.

121

A.J. Bruce and K.E. Coombcs

differs from both the other post-larvae in having
the second pereiopods already distinctly un¬
equal, and the fingers illustrated show the char¬
acteristic dactylar molar process, with an oppos¬
ing fossa on the fixed finger. The fingers of the
minor chela are not described or illustrated, but
may be presumed to lack these features. As the
specimen was obtained by moult from an earlier
larval stage, there is no doubt that it is the first
post-larval stage. As the larvae were obtained
from the plankton, the identity of the species is
uncertain, especially as some 11 species of
Periclimenaeus are now known from the tropi¬
cal western Atlantic Ocean.

A minor point of additional interest is that in
the specimens referred to Periclimenaeus
(?)wilsoni, the corpus of the ambulatory dactyl
bears a small acute tooth proximally. Such a
tooth is not present in adults of P. wilsoni. A
similar tooth may be found in the adults of
several Indo-West Pacific species that occur in
associations with compound ascidians. A simi¬
lar basal tooth is present on the ambulatory
dactyl of P. ascidiarum Holthuis, 1951, and it is
possible that these post-larvae should be re¬
ferred to this or some related species. It is
noteworthy that this apomorphic feature may be
present at such an early stage. It may also be
noted that Gurney and Lebour (1941) describe
and illustrate the posterior margin of the telson
as presenting “two pairs of apical spines, with a
pair of feathered setae between them”, suggest¬
ing a close relationship to shrimps of the sub¬
family Palaemoninae. However, in the present
specimen, the submedian spines, although
setulose, are robust and rigid, and may appropri¬
ately be considered to be “spines”, and thus to
conform at this early ontogenic stage, to the
current definition of the Pontoniinae.

The present post-larval specimen is provi¬
sionally referred to P. stylirostris on account of
the highly characteristic fingers of the second
pereiopods which are virtually identical with
those of the minor chela of adult P. stylirostris as
figured in Bmce (1972a). This species has not
been formally recorded in Australian waters but
the specimen reported from Elizabeth Reef, South
Pacific Ocean ( Periclimenaeus in Lowry and
Springthorpe 1992) has been re-studied and
found to be an ovigerous female of this species.
The fingers of the minor second pereiopod are
exactly as in the juvenile specimen, but the
rostrum has a dentition of 7/0. The palms of both
chelae are smooth and without denticles.

Australian distribution. Elizabeth Reef,
Coral Sea. Northern Territory: not previously
recorded.

Further distribution. Type locality: South
China Sea, 20°34.0’N, 113°30.5’E, ca. 90m.
Also known from Viti Levu, Fijian Islands.

Periclimenaeus tridentatus (Miers)

Coralliocaris tridentatus Miers, 1884: 2946,
pi. 32C.

Periclimenaeus tridentatus - Holthuis 1952:
140-146, figs 63-65 (partimf, - Bruce 1983a: 44-
45; 1993c: 834-838.

Material examined. Id", I 9 , stn CP/ 6 , Coral
Bay, 3-4 m, 23 June 1981, coll. A.J. Bruce et al.,
in unidentified colonial ascidian, NTM
Cr.000097. 1 ovig. 9 , same data as previous,
NTM Cr.000150. 1 ovig. 9 . same data as previ¬
ous, NTM Cr.008564. lo\ l9,stnCP/16,Trepang
Bay, 2 m, 14 October 1981, coll. A.J. Bruce et
al., in unidentified ascidian, NTM Cr.001871.
2d’, 29, 1 ovig. 9 , stn CP/21, Coral Bay, 18
October 1981, coll. A.J. Bmce et al., NTM
Cr.000156. 2d", 2 ovig. 9 , same data as previous,
NTM Cr.000157. Id", stn NY/4, McCluer Is¬
land, 7 m, 16 October 1982, coll. A.J. Bmce, in
unidentified ascidian. NTM Cr.009313.

Remarks. The specimens NTM Cr.000097
and NTM Cr.008564 have been reported upon
by Bmce 1993c.

Australian distribution. Type locality: Thurs¬
day Island, Torres Straits. Northern Territory:
Port Essington (Bmce 1983a). Western Aus¬
tralia: Cape Jaubert. Queensland: Heron Island,
Wistari Reef.

Further distribution. Also reported from
Mozambique; Singapore; Indonesia; Mariana
Islands; Johnson and Palmyra Islands, and Pearl
and Hermes Reefs.

Periclimenella spinifera (De Man)

Periclimenes Petitthouarsi var. spinifera De
Man, 1902: 284.

Periclimenes (Ancylocaris) spiniferus - Kemp
1922: 195-196.

Periclimenes (Harpilius) spiniferus - Holthuis
1952: 76-78, fig. 30.

Periclimenes spiniferus - Bmce 1983a: 42;
1988b: 229.

Periclimenella spinifera - Duris and Bmce
1995:656-661, figs 19-21.

122

The palaemonoid shrimp fauna of the Cobourg Peninsula

Material examined. 2d", 1 ovig. 9 , stn CP/2,
Kennedy Bay. 2 m, 22 June 1981, coll. A.J.
Bruce, P. Horner, J.N.A. Hooper, on coral
Acropora sp., NTM Cr.007618. 1 ovig. 9 , same
data as previous, NTMCr.008544. lo", 1 ovig. 9 ,
stn CP/5, Coral Bay, 2-3 m, 23 June 1981, coll.
A.J. Bruce, P. Horner, J.N.A. Hooper, on
Acropora sp., NTM Cr.008548. 1 spm., same
data as previous, NTM Cr.008552. lcf, 1 ovig. 9 ,
same data as previous, NTM Cr.008559. Id 1 ,
same data as previous, NTM Cr.008560.2 spms,
stn CP/16,Trepang Bay, 2 m, 14 October 1981,
coll. A. Bruce et al., on coral Seriatopora sp.,
NTM Cr.001903. 3 spms (1 ovig. 9 ), same data
as previous, on coral Stylophora sp., NTM
Cr.008643.1 ovig. 9 , stn NY/5, McCluer Island,
1 m, 16 October 1982, coll. A.J. Bruce, P.
Homer, NTM Cr.009319. 2 ovig. 9 , stn NY/ 8 ,
Oxley Island. LWS, 19 October 1982, coll. J.
Robinson, A.J. Bruce, NTM Cr.007764.7 spms
(3 ovig. 9 , 1 juv.), stn NY/9, Oxley Island, LWS,
20 October 1982, coll. A.J. Bruce, NTM
Cr.003155. Id", stn CP/64, Coral Bay, 3-4 m, 18
May 1983, coll. N.L. Bruce, on Acropora sp.,
NTM Cr.009274.1 spm., same data as previous,
NTM Cr.009345. 3 ovig. 9 , stn CP/65, Coral
Bay,0.5 m, 18May 1983, coll. N.L. Bruce,NTM
Cr.009282. Id", 2 ovig. 9 , stn CP/75, Coral Bay,
ca. 5 m, 15 September 1985, coll. J.E.N. Veron,
R. Williams, on Acropora samoensis, NTM
Cr.007570. 6 spms, stn CP/95, Coral Bay, 5-6
m, 12 August 1986, coll. R. Williams et al., on
Stylophora sp., NTM Cr.004154.

Australian distribution. Northern Territory:
Dudley Point, Darwin; Darwin Harbour (Bruce
1983a, 1987c). Queensland: Heron Island, One
Tree Island, Low Isles, North West Island.

Further distribution. Type locality: Temate,
Indonesia. Also reported from Kenya; Tangan¬
yika; Madagascar; Seychelle Islands; La
Reunion; Gulf of Manaar, India; Maidive Is¬
lands; Lakshadweep; Chagos Islands; Nicobar
Islands; Mergui Islands; Perhentian Islands;
Singapore; South China Sea; Philippines; Indo¬
nesia; Papua New Guinea; Ryukyu Islands;
Marshall Islands; Marianas Islands; Fiji Is¬
lands; Samoa; Wake Island and Tahiti.

Periclimenes affinis (Zehntner)

Palaemonella affinis Zehntner, 1894: 208.

Periclimenes (Harpilius) affinis - Holthuis
1958: 6 - 8 , fig. 2.

Material examined. 64 spms (16 ovig. 9 ), stn
CP/30, Black Point, 10-12 m, 29 April 1982,

coll. P. Horner, H.K. Larson, on crinoid
Heterometra magnipinna, NTM Cr.000294 A
& B.

Remarks. The specimens were all collected
from a single host crinoid, Heterometra
magnipinna A.H. Clarke (det. F. Rowe) and
represent an unusually large population of asso¬
ciated shrimps from a single host crinoid.
Periclimenes affinis has been previously associ¬
ated with the crinoids Comanthina schlegeli,
Comatula cratera and Comanthus sp. The asso¬
ciation with Heterometra represents a new host
record.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Wistari
Reef, Capricorn Islands.

Further distribution. Type locality: Ambon,
Indonesia. Also recorded from the South China
Sea; New Caledonia; Philippines; Indonesia and
(?) Japan.

Periclimenes alegrias Bruce

Periclimenes alegrias Bruce, 1987a: 143-
151, figs 1 a, 2-5,15 a-c; 1988b: 228; 1990b: 11.

Material examined. 1 ovig .9 paratype, stn
CP/70, Coral Bay, 2-5 m, 12 September 1985,
coll. L. Vail, on crinoid Lamprometra palmata,
NTM Cr.003223. 1 ovig .9 holotype, stn CP/76,
Coral Bay, 6 m, 15 September 1985, coll. L.
Vail, on crinoid Stephanometra spicata, NTM
Cr.004071. 10", 1 ovig .9 paratypes, stn CP/97,
Coral Bay, 12 August 1986, coll. A. Hoggett, on
crinoid Lamprometra klunzingeri, NTM
Cr.004168.3 juvs, stn CP/99, OrontesReef, 11-
12 m, 13 August 1986, coll. A. Hoggett, on
Lamprometra klunzingeri, NTM Cr.004173.

Australian distribution. Type locality: Port
Essington, Northern Territory. Northern Terri¬
tory: East Point, Darwin (Bruce 1988b); North
West Vernon Island (Bruce 1990b).

Further distribution. Not known outside
Australian waters.

Periclimenes amymone De Man

Periclimenes amymone De Man, 1902: 829-
833, pi. 25, fig. 53; - Bruce 1988b: 228.

Material examined, lo", 1 ovig. 9 , stn CP/2,
Kennedy Bay, 2 m, 22 June 1981, coll. A.J.
Bruce, J.N.A. Hooper, P. Horner, on coral
Acropora sp., NTM Cr.008531. 6 spms (2
ovig. 9 ), same data as previous, on Acropora
sp., NTM Cr.008537. Id", 1 ovig. 9 ,2 juv., same
data as previous, NTM Cr.008540. Id", 2 ovig. 9 ,

123

A.J. Bruce and K.E. Coombes

1 juv., stn CP/5, Coral Bay, 2-3 m, 23 June

1981, coll. A.J. Bruce, J.N.A. Hooper, P. Homer,
on Acropora sp„ NTM Cr.008536. Id", 1 ovig. 9 ,

2 juv., same data as previous, on Acropora sp.,
NTM Cr.008558. Id", 1 ovig. 9 , stn CP/ 6 , Coral
Bay, 3-4 m, 23 June 1981, coll. J.N.A. Hooper,
A.J. Bruce, P. Homer, on Acropora sp., NTM
Cr.008539. Id 1 , 1 ovig. 9 , same data as previ¬
ous, NTM Cr.008551. 1 spm., stn CP/13, Coral
Bay, 0.5 m, 20 July 1981, coll. J.R. Hanley et
al., on Acropora sp., NTM Cr.003007. 8 spms,
stn CP/16,Trepang Bay, 2 m, 14 October 1981,
coll. A.J. Bruce, J.N.A. Hooper, P. Homer, on
coral Seriatopora sp., NTM Cr.001902.20 spms
(7 ovig. 9 ), same data as previous, on coral
Stylophora sp., NTM Cr.008645. 12 spms, stn
CP/20, Walford Point, Coral Bay, LWS, 17
October 1981, coll. A.J. Bruce et al„ NTM
Cr.001948. 8 spms, stn CP/21, Coral Bay, 1-3
m, 18 October 1981. coll. P.N. Alderslade et
al., NTM Cr.001881. 1 spm., stn CP/33, Port
Bremer, 6 m, 1 May 1982, coll. J.N.A. Hooper,
P.N. Alderslade, NTM Cr.009331. 4 spms, stn
CP/37, Table Head, 1-3 m, 3 May 1982, coll. P.
Horner, on Acropora sp., NTM Cr.000304. 1
spm., stn NY/1, New Year Island, 10 m. 13
October 1982, coll. P. Homer, on Seriatopora
sp., NTM Cr.003174. 1 spm., same data as
previous, on Seriatopora sp., NTM Cr.007755.
8 spms (5 ovig. 9 ), same data as previous, on
Seriatopora sp., NTM Cr.007757. Id", 4 ovig. 9 ,
stn NY/2, New Year Island, 10 m, 14 October

1982, coll. A.J. Bruce, on Seriatopora sp.. NTM
Cr.009290.5 spms, same data as previous, NTM
Cr.009293. 4 ovig. 9 , stn NY/3, New Year Is¬
land, 16 m, 14 October 1982, coll. P. Homer et
al., NTM Cr.007777. Id 1 , 2 ovig. 9 , stn NY/4,
McCluer Island, 7 m, 16 October 1982, coll.
A.J. Bruce, NTM Cr.009277. 2o\ 1 ovig. 9 , I 9 ,
stn NY/5, McCluer Island, 1 m, 16 October
1982, coll. A.J. Bruce, P. Horner. NTM
Cr.009323. 1 ovig. 9 , I 9 . same data as previous,
NTM Cr.009325. 2 spms, same data as previ¬
ous, NTM Cr.009330. 4 spms (1 ovig. 9 ), stn
NY/9, Oxley Island, LWS, 20 October 1982,
coll. A.J. Bruce, NTM Cr.009292. 18 spms (2
ovig. 9 ), same data as previous, NTM Cr.009327.
6 spms, stn CP/51, Table Head, 13 May 1983,
coll. N.L. Bruce, on Acropora sp., NTM
Cr.010248. 2 spms, stn CP/60, Coial Bay, <6
m, 16 May 1983, coll. N.L. Brace, D. Staples,
on Acropora sp., bopyridized, NTM Cr.010130.

3 spms, same data as previous, on Stylophora
sp., NTM Cr.010251. 3 spms (1 juv.

bopyridized), same data as previous, on
Stylophora sp., NTM Cr.010517. 2 spms, stn
CP/61, Coral Bay, 4 m, 17 May 1983, coll. N.L.
Brace, J.R. Hanley, NTM Cr.010240. 3 spms,
same data as previous, NTM Cr.010246.7 spms,
same data as previous, on Acropora sp., NTM
Cr.010250.7 spms, same data as previous, NTM
Cr.010518. 8 spms (1 ovig. 9 ), stn CP/64,
Walford Point, 3-4 m, 18 May 1983, coll. N.L.
Bruce, NTM Cr.009338. 1 ovig. 9 , same data as
previous, NTM Cr.010243. 2 spms, same data
as previous, on Acropora sp., NTM Cr.010519.
lo", stn CP/69, Table Head, 1-4 m. 12 Septem¬
ber 1985, coll. J.E.N. Veron, J.N.A. Hooper, on
Acropora samoensis, NTM Cr.007622. 1 spm.,
same data as previous, on Acropora subulata,
NTM Cr.007623. 5 spms (1 ovig. 9 ), stn CP/70,
Coral Bay, 2-5 m, 12 September 1985, coll.
A.J. Bruce, on Stylophora (pistillatal). NTM
Cr.007556. 3 spms (1 ovig. 9 ), stn CP/71, Coral
Bay, 2-5 m, 13 September 1985, coll. A.J. Brace,
on Stylophora {pistillatal ), NTM Cr.007562. 1
ovig. 9 , same data as previous, on Stylophora
pistillata, NTM Cr. 007563. lo", 2 ovig. 9 , same
data as previous, on Stylophora sp., NTM
Cr.007564. 1 ovig. 9 , same data as previous, on
Acropora subulata, NTM Cr.007566. Id", 2
ovig. 9 , stn CP/75, Coral Bay, ca. 5 m, 15 Sep¬
tember 1985, coll. J.E.N. Veron, R. Williams,
on Acropora samoensis, NTM Cr.007571. 16
d", I 9 , same data as previous, on Stylophora
pistillata, NTM Cr.007572. 13 spms (5 ovig. 9 ,
3 juv.) same data as previous, on Acropora
latistella, NTM Cr.007573. 1 ovig. 9 , 19,3 juv.,
same data as previous, on Acropora samoensis,
NTM Cr.007604. 3 ovig. 9 , same data as previ¬
ous, on Stylophora sp., NTM Cr.007640. 1
ovig. 9 , stn CP/76, Coral Bay, 6 m, 15 Septem¬
ber, 1985, coll. L. Vail, J.E.N. Veron, R.
Williams, on Stylophora pistillata, NTM
Cr.007552. lo", same data as previous, on
Acropora latistella, NTM Cr.007635. 2 spms
(9 with hemiarthrinid bopyrid), stn CP/82, Coral
Bay, 5-6 m, 18 September 1985, coll. P.N.
Alderslade, R. Williams, on Acropora sp., NTM
Cr.007576. 6 spms (3 ovig. 9 ), same data as
previous, on Acropora digitifera , NTM
Cr.007606. 2o", 1 ovig. 9 , 4 juv., same data as
previous, on Stylophora pistillata, NTM
Cr.007610. 3 juv., stn CP/ 88 , Orontes Reef, 8 -
10 m, 9 August 1986, coll. C. Johnson, R.C.
Willan, on Acropora sp., NTM Cr.004102. 5
spms, stn CP/91, Orontes Reef, 10 August 1986,
coll. S. Slack-Smith, on coral Pocillopora sp..

124

The palaemonoid shrimp fauna of the Cobourg Peninsula

NTM Cr.004119. 3 spms, stn CP/93, Coral
Bay, 6-8 m, 11 August 1986, coll. R. Williams
et al., NTM Cr.004132. 4 spms, CP/95, Coral
Bay, 5-6 m, 12 August 1986, coll. R. Williams
et al., on Stylophora sp., NTM Cr.004156.

Australian distribution. Northern Territory:
Dudley Point, Darwin (Bruce 1983a, 1988b).
Queensland: Coral Sea, Heron Island, One Tree
Island.

Further distribution. Type locality: Ternate,
Indonesia. Also known from Nicobar Islands;
Singapore; Indonesia; New Caledonia; Papua-
New Guinea and Philippines.

Periclimenes anacanthus Bruce
(Fig. 12a)

Periclimenes anacanthus Bruce, 1988a: 105-
114, figs 1-5.

Material examined. 1 ovig. 9 , stn CPV/ 8 ,
West of Barrow Bay, LWS, 18September 1985,
coll. J.R. Hanley et al., NTM Cr.007652.

Remarks. The single example agrees well
with the original description and is only the
second record of this species. The rostrum has a
dentition of 1 + 8/3 and the merus of the single
second pereiopod is without any trace of a
distoventral tooth. The fingers have a small
proximal diastema with small acute teeth on
each side (Fig. 12a).

Australian distribution. Type locality:
Dunwich, North Stradbroke Island, Queens¬
land. Northern Territory: not previously re¬
corded.

Further distribution. Not recorded outside
Australian waters.

Periclimenes brevicarpalis (Schenkel)

Ancylocaris brevicarpalis Schenkel, 1902:
563, pi. 13, fig. 21; - Bruce 1988b: 228.

Periclimenes (Ancylocaris) brevicarpalis -
Kemp 1922: 185-191, figs 40-42, pi. 67.

Periclimenes (Harpilius) brevicarpalis -
Holthuis 1952: 69-73, fig. 27.

Periclimenes brevicarpalis - Bruce 1983a: 43.

Material examined, lcf, 1 ovig. 9 , stn CP/ 68 ,
Coral Bay, 2-3 m, 19 May 1983, coll. N.L.
Bruce, J.R. Hanley, from unidentified anemone,
NTM Cr.000442. 1 spm., stn CP/97, Coral Bay,
12 August 1986, coll. C. Johnson, from uniden¬
tified anemone, NTM Cr.004165.

Australian distribution. Northern Territory:
Dudley Point, Darwin (Bruce 1983a, 1988b).

Queensland: Heron Island, Magnetic Island,
Murray Island, Torres Strait, Low Isles, Saddle¬
back Island, Hope Island, Port Denison. West¬
ern Australia: Hermite Island, Monte Bello Is¬
lands.

Further distribution. Type locality: Makas¬
sar, Sulawesi, Indonesia. Also known from the
Red Sea; throughout most of the Indian Ocean,
and western Pacific to Japan; Solomon Islands,
Vanuatu; Caroline Islands and Marshall Is¬
lands.

Periclimenes cobourgi sp. nov.

(Figs 10-11)

Type material. 1 ovig .9 holotype, stn CP/37,
Table Head, 3 m, 3 May 1982, coll. H.K. Larson,
P. Homer, on gorgonian Rumphella aggregata,
NTM Cr.000307.

Diagnosis. A small sized species of Periclimenes,
of slender, subcylindrical body form.

Rostrum (Fig. 11a) well developed, acute,
reaching almost to distal margin of intermediate
segment of antennular peduncle, horizontal,
dorsal carina well developed, convex, with seven
acute teeth, all anterior to posterior orbital mar¬
gin, postrostral carina extending over anterior
halfof carapace; lateral carinae obsolete, ventral
carina absent, lower margin feebly biconvex,
distally unarmed, with few short setae proxi¬
mal ly.

Carapace (Fig. 10a) smooth, with well devel¬
oped epigastric spine at about 0.3 of carapace
length, on postrostral carina; inferior orbital
angle produced,distally rounded,antennal spine
acute, marginal, hepatic spine at about 0.2 of
carapace length, anterolateral angle of
branchiostegite bluntly obtuse.

Abdominal segments normal, third segment
not posterodorsally produced, sixth segment
twice the fifth segment length, twice as long
than deep. Telson (Fig. 11 m) about 0.9 of sixth
segment length, 3.75 times longer than anterior
of telson width, posterior margin (Fig. 1 In)
aboutO.33 of anterior width, angular, with small
acute median process, lateral spines small,
slightly larger than dorsal spines, intermediate
spines about 0.15 of telson length, submedian
spines about 0.5 of intermediate spine length,
setulose.

Antenna (Fig. 1 lb) with proximal segment of
peduncle about 2.6 times longer than wide,
anterolateral margin produced, rounded lobe
medially with small acute tooth laterally, lateral

125

A.J. Bruce and K.E. Coombes

margin convex, sty locerite slender, acute, reach¬
ing 0.5 of segment length, statocyst normally
developed, ventromedial margin withsmall acute
tooth; intermediate and distal segments com¬
bined length about 0.45 of proximal segment
length; upper ramus with four fused segments,
shorter ramus with single free segment, with six
groups of aesthetascs, longer ramus filiform;
lower ramus filiform.

Antenna (Fig. 1 lc) with basicerite with acute
distoventral tooth, carpocerite reaching to about
middle of scaphocerite; scaphocerite about 3.8
times longer than broad, with strong distolateral
tooth at about 0.75 of length, far exceeding
antennular peduncle.

Eye (Fig. lid) well developed with large
globular cornea, diameter about 0.27 of cara¬
pace length, peduncle elongate, about 1.6 times
longer than proximal width, tapering slightly
distally, accessory pigment spot well developed,
on raised tubercle (Fig. 1 le); eye length about
0.6 of carapace length.

Third maxilliped (Fig. Ilf) normal,
ischiomerus feebly separatedfrom basis, exopod
with four plumose setae distally, epipod large,
subcircular, without arthrobranch.

Thoracic sternites broad, fourth stemite with
stout anterior median process (Fig. 1 lg)

First pereiopods (Fig. 1 lg) extend to end of
antennularpeduncle. Chela (Fig. 1 lh) with palm
2.0 times longer than deep, compressed, fingers
subequal to palm length, slender, with acute,
feebly hooked simple tips, cutting edges entire,
lateral, extending over distal 0.6 of fingers;
carpus subequal tochela;merus 1.1 times carpus
length, subcylindrical,unarmed; coxa with small
distoventral lobe.

Second pereiopods (Fig. 1 li) feeble, subequal,
similar, reaching to end of scaphocerite; chela
(Fig. 1 lj) small, about 0.55 of carapace length,
palm 2.0 times longer than deep, slightly swol¬
len, compressed, smooth, fingers subequal to
palm length, slender, with lateral entire cutting
edges over distal 0.6, tips simple, feebly hooked;
carpus about 0.9 of chela length, unarmed, slen¬
der, 8.0 times longer than sub-distal width;
merus 0.8 of carpal length, subcylindrical, 7.0
times longer than wide, unarmed; ischium 1.2
times meral length.

Ambulatory pereiopods slender. Third
pereiopod with dactyl (Fig. 111) about 0.33 of
propod length, slender, compressed, unguis not
distinctly demarcated, about 0.8 of corpus
length, compressed, with sharp ventral margin,
accessory tooth well developed, 0.65 of unguis
length; propod (Fig. 1 lk) 10.0 times longer

Fig. 10. A, Periclimenes cobourgi sp. nov., holotype female. Table Head, Port Essington; B, Periclimenes indicus (Kemp),
ovigenous female (after Kemp (1915)).

126

The palaemonoid shrimp fauna of the Cobourg Peninsula

than deep, with one pair of long distoventral
spines, longer spine about 0.45 of dactyl length,
one pair of distal ventral spines, two long ven¬
tral spines.

Uropods normal, exceeding telson. Ova small,
ca. 0.5mm.

Measurement. Ovigerous female, carapace
length 1.5mm.

Fig. 11. Periclimenes cobourgi sp. nov., ovigerous female holotype. A, anterior carapace, rostrum; B, antennule; C, antenna;
D, eye, dorsal; E, same, posterior, F, third maxilliped; G, first pereiopod; H, same, chela; 1, second pereiopod, J, same, chela,
K, third pereiopod, propod, dactyl; L, same, dactyl; M, telson; N, same, posterior margin.

127

A.J. Bruce and K.E. Coombes

Table 2. Periclimenes cobourgi sp.nov. may be distinguished from P. indicus by the following characteristics.

Periclimenes cobourgi sp. nov.

1. Rostral dentition 7/0.

2. Dorsal margin of rostrum convex, rostrum not appearing
angular.

3. Rostrum not exceeding intermediate segment of antennular
peduncle.

4. Epigastric spine at about 0.3 of CL.

5. Hepatic spine at lower, more anterior position (Fig. 9a).

6. Upper flagellum of antennule with first four segments
fused, one free segment.

7. Scaphocerite far exceeding antennular peduncle.

8. Eye stalk reaching 0.8 of rostral length, greater than 0.6
of CL.

9. Anterolateral angle of branchiostegite bluntly obtuse.

10. Comeal diameter about 0.25 of CL, 0.65 of stalk length,
with accessory pigment spot on raised tubercle.

11. Third ambulatory pereiopod with long ventral spines over
distal half of propod: spines long, greater than propod
width, about half dactyl length.

12. Ambulatory dactyl with accessory tooth about 0.6 of
length of unguis.

13. Telson posterior margin with small acute median process.

Periclimenes indicus (Kemp)

Rostral dentition 8-10/1-3

Dorsal margin of rostrum sublinear, giving rostrum an angu¬
lar shape.

Rostrum distinctly exceeding intermediate segment of
antennular peduncle.

Epigastric spine at about 0.5 of CL.

Hepatic spine at higher, more posterior position (Fig. 9b).

Upper antennular flagellum with first nine segments fused,
two to three free segments.

Scaphocerite only slightly exceeding antennular peduncle.
Eye stalk reaching about 0.6 of rostral length, distinctly less
than 0.5 of CL.

Anterolateral angle of branchiostegite bluntly subreetanglar,
feebly produced.

Comeal diameter about 0.22 of CL, 0.87 of stalk length,
accessory pigment spot not on raised tubercle.

Third ambulatory pereiopod with 7-8 spines over whole sev¬
eral propod length: spines short, less than propod width, much
less than half dactyl length.

Ambulatory dactyl with accessory tooth less than half length
of unguis.

Telson posterior margin without acute median process.

Host. Rumphella aggregata (Nutting)
(Gorgonacea).

Etymology. The specific name is derived
from the collection locality, the Cobourg Penin¬
sula, Arnhem Land.

Systematic position. Periclimenes cobourgi
appears closely related only to Periclimenes
indicus (Kemp, 1915), a species that shows
some resemblance to the Periclimenes grandis
species group sensu lato (Bruce 1987d), on
account of the presence of a median process on
the anterior fourth thoracic stemite. Both spe¬
cies are unusual in that group in having strongly
biunguiculate dactyls on the ambulatory
pereiopods, and also differ significantly from
most species of the group in the very feeble
development of the second pereiopods. They
appear to occupy a rather isolated systematic
position among the numerous species of the
genus Periclimenes, and are intermediate be¬
tween the “ grandis ” - group species sensu lato
and those of the rest of the genus Periclimenes
sensu stricto.

Remarks. The association of P. cobourgi
with a gorgonian host contrasts strongly with
the habits of P. indicus. P. indicus was first
reported from Chilka Lake, a brackish water

situation (Kemp 1915). The validity of the as¬
sociation with this gorgonian host is also sup¬
ported by the presence of broad thoracic
sternites, a common feature in other gorgonian
associated genera such as Pontonides and
Hamodactylus.

Periclimenes commensalis Borradaile

Periclimenes (Cristiger) commensalis
Borradaile, 1915: 211.

Periclimenes (Periclimenes) commensalis -
Clark 1921: 628.

Allopontonia iaini - Lowry and Springthorpe
1992: 128.

Material examined. 1 spm.,stn CP/23, Walford
Point, 5 m, 19 October 1981, NTM Cr.005351. 1
spm., stn CP/24, Smith Point, 4-5 m, 19 October
1981, NTM Cr.005359.1 9 , same data as previous,
NTM Cr.005362.1 spm., stn CP/26, Sandy Island
No. 2,7 m,20October 1981, coll. P.N. Alderslade,
J.N.A. Hooper, on soft coral Telesio sp., NTM
Cr.005356. 1 ovig. 9 , stn NY/2, New Year Island,
10 m, 14 October 1982, coll. A.J. Bruce, NTM
Cr.009296. lo\ same data as previous, on uniden¬
tified crinoid, NTM Cr.009297. lo\ 1 ovig. 9 , same
data as previous, NTM Cr.009299.

128

The palaemonoid shrimp fauna of the Cobourg Peninsula

Remarks. This species is a well known as¬
sociate of crinoid echinoderms. The specimens
referred to Allopontonia iaini from Elizabeth
and Middleton Reefs, Coral Sea (Lowry and
Springthorpe 1992) have been re-examined and
their material from Oxycomanthus bennetti was
found to belong to Periclimenes commensalis.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island, Bribie Passage, Myora. North Stradbroke
Island, Elizabeth and Middleton Reefs.

Further distribution. Type locality: Murray
Island, Torres Strait, Queensland. Also known
from Zanzibar; Kenya; Mozambique; Indone¬
sia; Philippines; Hong Kong; Japan; Ryukyu
Islands; Marshal 1 Islands; New Caledonia; Solo¬
mon Islands and Fijian Islands.

Periclimenes diversipes Kemp

Periclimenes (Ancylocaris)diversipes Kemp,
1922: 179-184 (partim), figs 36-39; - Bruce
1987c: 37.

Material examined. 1 ovig. 9 , stn CP/2,
Kennedy Bay,2 m, 22 June 1981, coll. A.J. Bruce,
J.N.A. Hooper, P. Homer, on coral Turbinaria
sp., NTM Cr.008565. 1 ovig. 9 , stn CP/5, Coral
Bay, 2-3 m, 23 June 1981, coll. A.J. Bruce, J.N.A.
Hooper, P. Horner, on Turbinaria sp., NTM
Cr.008554. Id", 2 ovig. 9 , same data as previous,
on coral Acropora sp., NTM Cr.008561.5 spms
( 2 ovig. 9 ), stn CP/61, Coral Bay, 4 m, 17 May
1983, coll. N.L. Bruce, J.R. Hanley, on coral
Herpolitha Umax, NTM Cr.000438.

Remarks. The association of this common
coral commensal with Herpolitha represents a
new host record.

Australian distribution. Northern Territory:
Darwin Harbour (Bruce 1987c). Queensland:
Heron Island, Restoration Rock.

Further distribution. Type locality: Kilikarai,
Gulf of Manaar. Also reported from the Red Sea;
Aden; Kenya; Tanganyika; Comoro Islands;
Madagascar; Seychelle Islands; Zanzibar; La
Reunion; Singapore and Thailand.

Periclimenes elegans (Paulson)

Anchistia elegans Paulson, 1875: 113, pi. 17,
fig. 1 .

Periclimenes (Ancylocaris) elegans - Kemp
1922: 215-218, fig. 60-62.

Periclimenes (Harpilius) elegans - Holthuis
1952:81-82, fig. 31.

Periclimenes elegans - Bruce 1988b: 288.

Material examined. 1 spm., stn CP/2,
Kennedy Bay, 2 m, 22 June 1981, coll. J.N.A.
Hooper, P. Homer, A.J. Bruce, NTM Cr.002719.
3 spms, stn NY/ 8 , Oxley Island, LWS, 19 Octo¬
ber 1982, coll. J. Robinson, A.J. Bruce, NTM
Cr.007761.8 spms (4 ovig. 9 ,2 juv.), same data
as previous, NTM Cr.007762. 26 spms, stn NY/
9,Oxley Island, 20October 1982, coll. A.J. Bruce,
NTM Cr.003156.7 spms (3 ovig. 9 ), stn CP/45,
Table Head, LW, 11 May 1983, coll. A.J. Bruce,
NTM Cr.009346. 2<f, 1 ovig. 9 , stn CP/77,
Orontes Bay, 10 m, 16 September 1985, coll. R.
Williams, on coral Stylophora pistillata, NTM
Cr.007557.1 ovig. 9 , stn CP/85,Orontes Reef, 10-
15 m, 19 September 1985, coll. L. Vail, R.
Williams, C. Hood, on soft coral Dendronepthya
sp., NTM Cr.007615.

Remarks. The associations with Stylophora
and Dendronepthya are probably incidental or
accidental.

Australian distribution. Northern Territory:
East Point, Darwin (Bruce 1988b).

Further distribution. T ype locality: Red Sea.
Also recorded from Persian Gulf; Aden; Kenya;
Zanzibar; Tanganyika; Pakistan; Seychelle Is¬
lands; Nicobar Islands; Madagascar; Andaman
Islands; Sri Lanka; Ryukyu Islands; Hong Kong;
Indonesia; Singapore; Caroline Islands and
Marshall Islands.

Periclimenes galene Holthuis

Periclimenes (Harpilius) galene Holthuis,
1952: 11,62, fig. 24.

Periclimenes galene -Bmce 1976: 12, figs 3,
4; 1983c: 207.

Material examined. 1 ovig. 9 (CL 3.3mm),
stn NY/4, McCluer Island, 11°02.0’S 132°58.5
E, 7 m, 16 October 1982, on unidentified hydroid,
NTM CrO 11328.

Remarks. The specimen agrees exactly with the
original descriptions, with a slender rostrum with
a dentition of 1 + 5 + 1 + 1/0. The specimen was
largely transparent in life with the body with about
six fine longitudinal brown striae, first and second
pereiopods, antennal peduncles and caudal fan
speckled with brown dots, pleopods also striate.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island, Lizard Island.

Further distribution. Type locality: Menado,
Indonesia. Also known from Kenya; Zanzibar;
Tanganyika and Comoro islands.

129

A J. Bruce and K.b. Coombes

Periclimenes grandis (Stimpson)

Anchistia grandis Stimpson, 1860: 39.
Periclimenes grandis - Borradaile 1898: 382;

- Bruce 1987c: 37; 1988b: 229.

Material examined. 2 juv., stn CP/9, Coral
Bay, 0.5 m, 24 June 1981, coll. A.J. Bruce, on
coral Acropora sp., NTM Cr.008547. 4 spms (2
ovig. 9 ), stn CP/12, Coral Bay, 0.1 -0.5 m, 19 July

1981, coll. A.J. Bruce, NTM Cr.000122.1 ovig. 9 ,
stn CP/17, Wanaray Point, Trepang Bay, LWS,

15 October 1981, coll. A.J. Bruce, P. Homer,
NTM Cr.008696. 28 spms (5 ovig. 9 ), same data
as previous, NTM Cr.008715. 8 spms, stn CP/
18, Midjari Reef, Trepang Bay, LWS, 16 Octo¬
ber 1981, coll. A.J. Bruce e/a/., NTM Cr.008242.
2 spms, stn CP/38, Table Head, 2-4 m, 4 May

1982, coll. H.K. Larson etal. , NTMCr.001967.
lcf, stn NY/5, McCluer Island, 1 m, 16 October
1982, coll. A.J. Bruce, P. Horner, NTM
Cr.009321.1 ovig. 9 , stnCP/45,TableHead,LW,

11 May 1983, coll. A.J. Bruce, NTM Cr.009360.

Australian distribution. Northern Territory:
East Point, Darwin (Bruce 1987c, 1988b).
Queensland: Magnetic Island, PortMolle (?).

Further distribution. Type locality: Oshima,
Japan. Also recorded from the Red Sea; Jibouti;
Aden; Kenya; Zanzibar; Tanganyika; Madagas¬
car; Mo?ambique; Comoro Islands; Seychelle
Islands; India; Sri Lanka; Mergui Islands; Ma¬
laya; Singapore; Indonesia; China; Japan; Fi¬
jian Islands and Marshall Islands.

Periclimenes holthuisi Bruce

Periclimenes holthuisiBruce, 1969:258-259;
1982:244-246, fig. 7;-Bruce 1983a: 43; 1988b:
229.

Material examined. 1 spm., stn CP/20, Coral
Bay, LWS, 17 October 1981, coll. A.J. Bruce et
al., NTM Cr.002663. 10 spms, stn CP/21 Coral
Bay, 1-3 m, 18 October 1981, coll. P.N.
Aldersladeefa/.,NTMCr.001885. lo", 1 ovig. 9 ,
stn CP/28, Sandy Island No. 2,6-7 m, 22 Octo¬
ber 1981, coll. J.R. Hanley, NTM Cr.000117. 1
spm., stn CP/35, Danger Point, Port Bremer, 4
m, 1 May 1982. coll. A.J. Bruce, NTM
Cr.001730.4spms, stn CP/60, Coral Bay, <6 m,

16 May 1983, coll. N.L. Bruce, D. Staples, on
coral Heliofungia sp., NTM Cr.010247. 1 juv.,
stn CP/ 68 , Coral Bay, 2-3 m, 19 May 1983, coll.
J.R. Hanley, N.L. Bruce, on unidentified
anemone, NTM Cr.010480. 1 juv., stn CP/75,
Coral Bay, ca. 5 m, 15 September 1985, coll.

J.E.N. Veron, R. Williams, on unidentified
anemone, NTM Cr.003231.3 spms (1 ovig. 9 ),
same data as previous, on coral Euphyllia sp.,
NTM Cr.003232.1 spm., same data as previous,
NTM Cr.006339. I 9 , stn CP/80, Orontes Reef,
10 m, 17 September 1985, coll. J.E.N. Veron, R.
Williams, NTM Cr.006344. 12 spms (8 ovig. 9 ),
stn CP/ 86 , Coral Bay, 8 m, 7 August 1986, coll.
C. Johnson et al., on unidentified anemone,
NTM Cr.004075. 4 spms, stn CP/99, Orontes
Reef, 11-12 m, 13 August 1986, coll. A. Hogget
etal., on unidentified anemone, NTM Cr.004175.

Remarks. The association of this common
coelenterate commensal with Heliofungia rep¬
resents a new host record.

Australian distribution. Northern Territory:
Cobourg Peninsula, Sandy Island No. 2 (Bruce
1983a) East Point, Darwin (Bruce 1988b).
Queensland: Morton Bay, Heron Island, Peloris
Island, Bowen.

Further distribution. Type locality: Lung Ha
Wan, Hong Kong. Also reported from the Red
Sea; Kenya; Zanzibar; Maidive Islands; Sri
Lanka; Malaya; Singapore; Indonesia; Papua
New Guinea; Japan; Philippines; New Caledo¬
nia; Lord Howe Island; Caroline Islands and
Marshall Islands.

Periclimenes incertus Borradaile

Periclimenes (Cristiger) incertus Borradaile,
1915: 210; 1917: 384, pi. 53, fig. 7.

Periclimenes incertus - Bruce 1988b: 229.

Material examined. 4spms, stn CP/27, Sandy
Island No. 2,10 m, 21 October 1981, coll. J.N. A.
Hooper et al., on sponge Ianthella basta, NTM
Cr.003206. 1 ovig. 9 , stn CP/28, Sandy Island
No. 2,6-7 m, 22 October 1981, coll. J.R. Hanley,
NTM Cr.000161. 1 ovig. 9 , stn CP/30, Black
Point, 10-12 m, 29 April 1982, coll. P. Horner,
H.K. Larson, NTM Cr.000295.1 ovig. 9 , stn CP/
44, Table Head, 4-5 m, 11 May 1983, coll. N.L.
Bruce, D. Staples, NTM Cr.009336. I 9 , same
data as previous, NTM Cr.009339. la", 1 ovig. 9 ,
stn CP/46, off Table Head, 5 m, 12 May 1983,
coll. N.L. Bruce, NTM Cr.009353. 2 spms (1
ovig. 9 ), stn CP/60, Coral Bay, <6 m, 16 May
1983, coll. N.L. Bruce, D. Staples, NTM
Cr.010520. 2a", 2 ovig. 9 , stn CP/62, Coral Bay,
4 m, 17 March 1983, coll. N.L. Bruce, J.R.
Hanley, NTM Cr.009343.9 spms (5 ovig. 9 ), stn
CP/73, Table Head, 6-8 m, 14 September 1985,
coll. P.N. Alderslade, R. Williams, from soft
coral Telesto sp., NTM Cr.004055. 1 ovig. 9 , stn

130

The palaemonoid shrimp fauna of the Cobourg Peninsula

CP/76, Coral Bay, 6 m, 15 September 1985,coll.
P.N. Alderslade, L. Vail, R. Williams, NTM
Cr.007626. lo", 1 ovig. 9 , 1 juv., stn CP/77,
Orontes Bay, 10 m, 16 September 1985, coll. R.
Williams, on coral Stylophora pistillata , NTM
Cr.007681.1 ovig. 9 , stn CPV/ 8 , West of Barrow
Bay,LWS, 18 September 1985, coll. J.R. Hanley
et al., NTM Cr.007650. 7 spms (2 ovig. 9 ), stn
CP/85, Orontes Reef, 10-15 m, 19 September
1985, coll. C. Hood etal., on Duncanopsammia
sp., NTM Cr.004057. 3 spms, same data as
previous, NTM Cr.007613. 2 spms, stn CP/ 86 ,
Table Head, 8 m. 7 August 1986, coll. C. Johnson
et al., on yellow sponge, NTM Cr.004076. 6
juvs, same dataas previous, on red sponge, NTM
Cr.004079. 1 juv., same data as previous, NTM
Cr.004093.2 juvs, stn CP/93, Coral Bay, 6-8 m,
11 August 1986,coll.R. Williams etal., on coral
Turbinaria sp., NTM Cr.004129. 25 spms
(mainly juveniles), same data as previous, on
bright red sponge, NTM Cr.004133. 3 ovig. 9 ,
same data as previous, on blue sponge, NTM
Cr.004143. 1 ovig. 9 , stn CP/99, Orontes Reef,
11-12 m, 13 August 1985, coll. G. Morgan, A.
Hoggett, C. Johnson, on colonial ascidian, NTM
Cr.004177.

Remarks. This species not been previously
reported in association with Telestacea. The
association with an ascidian may have been
accidental.

Australian distribution. Northern Territory:
East Point, Darwin (Bruce 1988b). Queensland:
Heron Island. Western Australia: North West
Cape.

Further distribution. Type locality: South
Nilandu Atoll, Maidive Islands. Also known
from Aden; Kenya; Zanzibar; Tanganyika;
Madagascar; Sri Lanka; Andaman Islands; Sin¬
gapore; Philippines and New Caledonia.

Periclimenes indicus (Kemp)

(Fig. 10b)

Urocaris indica Kemp, 1915: 275-279, fig.
26, pi. 13, fig. 9.

Periclimenes (Periclimenes) indicus - Kemp
1922: 144, fig. 13.

Material examined. 3 spms, stn CP/10, Black
Point, 1-2 m, 18 July 1981, coll. A.J. Bruce,
J.N.A. Hooper, from seaweed Sargassum sp.
washings, NTM Cr.001197. 12 spms, stn CP/
13, Coral Bay, 0.5 m, 20 July 1981, coll. A.J.
Bruce, J.N.A. Hooper, from Sargassum sp. wash¬
ings, NTM Cr.006496. 2 spms, stn CP/18,

Midjari Point, Trepang Bay, LWS, 16 October

1981, coll. A.J. Bruce et al., NTM Cr.008241.5
spms (2 ovig. 9 ), stn CP/58, Caiman Creek, <1
m, 15 May 1983, coll. N.L. Bruce, A.J. Bruce,
NTM Cr.009285.

Australian distribution. Northern Territory:
not previously reported. Queensland: Moreton
Bay.

Further distribution. Type locality: Chilka
Lake, India. Also known from Madras and Gulf
of Manaar, India; Nicobar Islands; Malaya;
Singapore and Sulawesi, Indonesia.

Periclimenes kempi Bruce

Periclimenes kempi Bruce, 1969: 260-261; -
Bruce 1988b: 229.

Material examined. I 9 , stn CP/2, Kennedy
Bay, 2 m, 22 June 1981, coll. A.J. Bruce, J.N.A.
Hooper, P. Homer, on soft coral Dendronephthya
sp., NTM Cr.008556. 1 spm., stn CP/5, Coral
Bay, 2-3 m, 23 June 1981, coll. A.J. Bruce,
J.N.A. Hooper, P. Homer, on coral Acropora
sp., NTM Cr.008535.1 ovig. 9 , same data as pre¬
vious, on yellow alcyonarian, NTM Cr.008555.
13 spms (1 ovig. 9 ) ( 2 s P ms with hemiarthrinid
bopyrids), stn CP/37, Table Head, 1 -3 m, 3 May

1982, coll. P.N. Alderslade, on soft coral Vepf/iya
sp., NTM Cr.000305. 1 spm., same data as
previous, coll. H.K. Larson, on soft coral
Stereonephthya sp., NTM Cr.002936. 2 spms,
stn CP/38, Table Head, 2-4 m, 4 May 1982, coll.
H.K. Larson, on soft coral Sinulariapolydactyla,
NTM Cr.001970.

Australian distribution. NorthemTerritory.
East Point, Darwin (Bruce 1988b). Queensland:
Broadhurst Reef, Heron Island.

Further distribution. Type locality:
Ghardaqa, Egypt. Also known from Kenya;
Zanzibar; Singapore; Philippines; Fijian and
Caroline Islands.

Periclimenes lutescens (Dana)

Harpilius lutescens Dana, 1852: 25.
Periclimenes (Ancylocaris) amamiensis -
Kubo 1940a: 44-46, figs 11-12.

Periclimenes (Harpilius) lutescens - Holthuis
1952: 88-91, fig. 35.

Material examined. 2 spms (1 ovig. 9 ), stn
CP/2, Kennedy Bay, 2 m, 22 June 1981, coll. P.
Homer, J.N.A. Hooper, A.J. Bruce, on coral
Acropora sp., NTM Cr.000096.5 spms, stn CP/
20, Walford Point, Coral Bay, LWS, 17 October

131

A.J. Bruce and K.E. Coombes

1981, coll. A.J. Bruce et al., on Acropora sp.,
NTM Cr.001947. lcT, stn NY/9, Oxley Island,
LWS, 20 October 1982, coll. A.J. Bruce, NTM
Cr.009316.2 spms, stn CP/51, Table Head, 4 m,
13 May 1983, coll. N.L. Bruce,on Acropora sp.,
NTM Cr.010482. 1 spm., stn CP/61, Coral Bay,
4m, 17May 1983, coll. N.L. Bruce, J.R. Hanley,
on Acropora sp., NTM Cr.010521. 1 ovig. 9 , stn
CP/69, Table Head, 1-4 in, 12 September 1985,
coll. J.E.N. Veron, J.N. A. Hooper, on Acropora
sp., NTM Cr.007551.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island, Capricorn Islands; Gillet Cay, Swain
Reefs.

Further distribution. Type locality: Tonga-
tabu, Cook Islands. Also known from the Red
Sea; Kenya; Zanzibar; Tanganyika; Comoro
Islands; Seychelle Islands; Madagascar;
Maidive Islands; Indonesia; Japan; Singapore;
Vietnam; Solomon Islands and Samoan Islands.

Periclimenes madreporae Bruce

Periclimenes madreporae Bruce, 1969: 262-
263.

Allopontonia iaini - Lowry and Springthorpe
1992: 128.

Material examined. 1 ovig. 9 , stn NY/ 8 , Oxley
Island, LWS, 19 October 1982, coll. A.J. Bruce,
on coral Goniopora sp., NTM Cr.006361.

Remarks. The specimens referred to
Allopontonia iaini from Elizabeth and Middleton
Reefs, Coral Sea (Lowry and Springthorpe 1992),
have been re-examined, with those specimens
from coral hosts found to belong to P.
madreporae.

Australian distribution. Type locality:
Erskine Island, Capricorn Islands, Queensland.
Northern Territory: not previously recorded.
Queensland: One Tree Island, Heron Island,
Willis Island, Bet Reef, Restoration Rock, Coral
Sea, Elizabeth and Middleton Reefs.

Further distribution. La Reunion; Solomon
Islands and Caroline Islands (?).

Periclimenes magnificus Bruce

Periclimenes magnificus Bruce, 1979a: 195-
207, figs 1-5, pis a-c.

Material examined. 2 spms, stn NY/10, Oxley
Island, 14 m, 21 October 1982, coll. A.J. Bruce et
al., on cerianthid anemone, NTM Cr.000387. lo\
same data as previous, NTM Cr.007726.2o\ I 9 ,1

juv., same data as previous, NTM Cr.007727. Id",
same data as previous, NTM Cr.007728. 1 spm.,
same data as previous, NTM Cr.007733. 1 spm.,
stn CP/78, Orontes Reef, 19 m, 16 September
1985, coll. R. Williams, on anemone Cerianthus
sp., NTM Cr.006346. 19 , stn CP/80, Orontes Reef,
10 m, 17 September 1985, coll. J.E.N. Veron, R.
Williams, on Cerianthus sp., NTM Cr.006343.

Australian distribution. Type locality:
Wistari Reef, Heron Island, Queensland. North¬
ern Territory: not previously recorded. Queens¬
land: Low Isles.

Further distribution. Reported also from the
Ryukyu Islands; Lesser Sunda Islands; Indone¬
sia and Cebu, Philippines.

Periclimenes mahei Bruce

Periclimenes mahei Bruce, 1969: 263-264.

Material examined. 1 spm., stn CP/5, Coral
Bay, 2-3 m, 23 June 1981, coll. A.J. Bruce,
J.N.A. Hooper, P. Homer, on coral Acropora
sp., NTM Cr.008550.

Australian distribution. Northern Territory:
not previously recorded. Western Australia: Point
Quobba.

Further distribution. Type locality: North¬
west Bay, Mahe, Seychelle Islands. Otherwise
recorded only from Zanzibar; Comoro Islands
and Amirante Islands.

Periclimenes nilandensis Borradaile

Periclimenes (Falciger) nilandensis
Borradaile, 1915:211; 1917:372, pi. 54, fig. 13.

Periclimenes (Harpilius) nilandensis -
Holthuis 1952: 58-60, fig. 22.

Material examined. 1 ovig. 9 , stn CP/14,
Burford Island, LWS, 13 October 1981, coll.
A.J. Bruce et al., NTM Cr.003191.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Wistari
Reef, Capricorn Islands.

Further distribution. Type locality: South
Nilandu Atoll, Maidive Islands. Also known
from Zanzibar; Kenya; Madagascar; Indonesia;
South China Sea and New Caledonia.

Periclimenes novaecaledoniae Bruce

Periclimenes novaecaledoniae Bruce, 1968:
1157-1165, figs 6-9.

Material examined, la", 1 ovig. 9 , stn CP/78,
Orontes Reef, 19 m, 16 September 1985, coll. L.

132

The palaemonoid shrimp fauna of the Cobourg Peninsula

Vail, on crinoid Lamprometra klunzingeri, NTM
Cr.007620.

Remarks. The association with the crinoid
Lamprometra klunzingeri represents a new host
record, as the type material was associated with
the crinoid Tropiometra afra.

Australian distribution. Northern Territory:
not previously recorded. New to Australian fauna.

Further distribution. Type locality: Ilot
Maitre, Noumea, New Caledonia. Otherwise
reported only from Madagascar (?).

Periclimenes obscurus Kemp

Periclimenes (Periclimenes)obscurus Kemp,
1922: 144-146, figs 14-15; - Bruce 1988b: 229.

Material examined. 1 spm.,stn CP/73, Table
Head, 6-8 m, 14 September 1985, coll. C. Hood
et a/., on soft coral Telesto sp., NTM Cr.007624.
1 ovig. 9 , stn CP/71, Orontes Bay, 10 m, 16
September 1985, coll. R. Williams, from coral
Seriatopora hystrix, NTM Cr.007630. 1 spm.,
stn CPV/ 8 , West of Barrow Bay, LWS, 18 Sep¬
tember 1985, coll J.R. Hanley et al., NTM
Cr.007647.

Australian distribution. Northern Territory:
East Point, Darwin (Bruce 1988b). Queensland:
Moreton Bay.

Further distribution. Type locality: Ennur
Backwater, Madras Harbour, India. Also re¬
corded from Kuwait; Kenya; Zanzibar; Tangan¬
yika and Madagascar.

Periclimenes seychellensis Borradaile

Periclimenes (Faleiger) seychellensis Bor¬
radaile, 1915:212; 1917: 374, pis. 55-55, fig. 14.

Periclimenes (Harpilius) seychellensis -
Holthuis 1952: 66-67, fig. 25.

Material examined. 1 spm., stn CP/10, Black
Point, 1-2 m, 18 July 1981, coll. A.J. Bruce,
J.N. A. Hooper, in seaweed Sargassum sp. wash¬
ings, NTM Cr.001 196.1 spm., stn CP/13, Coral
Bay, 0.5 m, 20 July 1981, coll. A.J. Bruce,
J.N.A. Hooper, NTM Cr.010264. 1 ovig. 9 , stn
CP/34, Danger Point, Port Bremer, 2 m, 1 May
1982, coll. J.R. Hanley, R. Williams, NTM
Cr.009334. 1 ovig. 9 , stn NY/ 8 , ° xle y Island ’
LWS, 19 October 1982, coll. J. Robinson, A.J.
Bruce, NTM Cr.007772.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island.

Further distribution. Type locality: Praslin,
Seychelle Islands. Also known from Kenya;

Zanzibar; Tanganyika; Mozambique; Madagas¬
car; Pakistan; South India; Singapore; Lesser
Sunda Islands; Indonesia and New Caledonia.

Periclimenes sinensis Bruce
(Figs 12b, c)

Periclimenes sinensis Bruce, 1969: 270-272;
1982: 255-258, figs 13-14.

Material examined. 1 ovig. 9 , stn NY/3, New
Year Island, 16 m, 14 October 1982, coll. P.
Homer etal., NTM Cr.007779. 1 ovig. 9 , stn CP/
43, Table Head, 2-4.5 m, 11 May 1983, on soft
coral Dendronephthya sp., NTM Cr.009660.

Remarks. The type specimens were found in
association with the alcyonarian Morchellana
planoregularis (Burchardt). The ovigerous fe¬
male (NTM Cr.009660) has a rostral dentition
of 9/2. The fourth thoracic sternite is broad with
a well developed transverse posterior ridge with
a pair of small acute submedian teeth separated
by a deep notch. A similar, less well developed
ridge is present on the fifth sternite. The basis
and coxa have only small setose ventromedial
lobes. The single second pereiopod is robust.
The dactyl has a single acute recurved tooth
proximally, the fixed finger has proximal series
of four teeth, the first three rounded and the
fourth acute, on a central cutting edge that
descends proximally into a fossa flanked by
medial and lateral edges.

Australian distribution. Not previously re¬
corded from Australian waters.

Further distribution. Type locality: Hong
Kong. Also known only from Japan and Philip¬
pines.

Periclimenes soror Nobili

Periclimenes soror Nobili, 1904: 232; - Bruce
1978a: 299-396, figs 1-6.

Periclimenes (Periclimenes) soror - Holthuis
1952:51-53, fig. 17.

Material examined. 1 ovig. 9 , stn NY/2, New
Year Island, 10 m, 14 October 1982, coll. A.J.
Bruce, on crown-of-thoms starfish Acanthaster
plane1 NTM Cr.009314. lo\ 2 ovig. 9 , stn NY/
3, New Year Island, 16 m, 140ctober 1982, coll.
P. Homer et al., on Acanthaster planci, NTM
Cr.007778.

Remarks. Both lots of specimens were ob¬
tained from Acanthaster planci (L.). Colour
patterns not recorded.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron

133

A.J. Bruce and K.E. Coombes

Island. New South Wales: Jervis Bay, Cutawong.
Western Australia: Exmouth (Gulf?), Dampier
Archipelago.

Further distribution. Type locality: Jibouti.
Also known from the Red Sea; Kenya; Zanzibar;

Tanganyika; Madagascar; Comoro Islands;
Seychelle Islands; Chagos Islands; Hong Kong;
Indonesia; Philippines; Sabah; Japan; Ryukyu
Islands; Caroline Islands; Marshall Islands;
Marianas Islands; Solomon Islands, New Cal-

Fig. 12. Periclimenes anacanthus Bruce, ovigerous female. A, chela of major second pereiopod. Periclimenes sinensis Bruce,
ovigerous female. B, anterior thoracic stemites; C. right second pereiopod. Periclimenes toloensis Bruce, ovigerous female. D,
carapace, rostrum and antennal peduncles; E, anterior carapace and rostrum; F, fourth thoracic stemite and first pereiopod, third
thoracic stemite shaded. Anchistioides willeyi (Borradaile), female. G, chela of second pereiopod; H, same, distal part of fixed
finger.

134

The palaemonoid shrimp fauna of the Cobourg Peninsula

edonia; Fijian Islands; Society Islands; Tuamotu
Islands and Hawaiian Islands. Also recorded in
the Eastern Pacific Region from the Gulf of
California, Panama and Colombia.

Periclimenes tenuipes Borradaile

Periclimenes tenuipes Borradaile, 1898:384;
- Bruce 1983a: 42-43; 1987a: 35; 1988b: 229.

Periclimenes (Ancylocaris) tenuipes - Kemp
1922: 195-196.

Periclimenes (Harpilius) tenuipes - Holthuis
1952: 84-85.

Material examined. lo\ 1 juv., stn CP/17,
Wanaray Point, Trepang Bay, LWS, 15 October
1981, coll. A.J. Bruce, P. Horner, NTM
Cr.008707. Id", same data as previous, NTM
Cr.008716. lcf, 1 ovig. 9 , same data as previous,
NTM Cr.008718. 1 ovig. 9 , stn CP/18, Midjari
Point, Trepang Bay, LWS, 16 October 1981,
coll. A.J. Bruce et al., NTMCr.007582. 2spms,
same data as previous, NTM Cr.007587.1 spm.,
same data as previous, NTM Cr.008243.1 spm.,
stn CP/20, Walford Point, Coral Bay, LWS, 17
October 1981, coll. A.J. Bruce et al., NTM
Cr.002662. 1 spm., stn CP/38, Table Head, 2-4
m, 4 May 1982, coll. J.R. Hanley et al., NTM
Cr.000258.2 spms, same data as previous, NTM
Cr.001961. 1 spm., stn CP/62, Coral Bay, 4 m,
17 May 1983, coll. N.L. Bruce, J.R. Hanley,
NTM Cr.010255.

Australian distribution. Northern Territory:
East Point, Darwin; Darwin Harbour (Bruce
1983a, 1988b). Queensland: Wistari Reef, Cap¬
ricorn Islands.

Further distribution. Type locality: Ralun,
New Britain. Also recorded from the Red Sea;
Kenya; Zanzibar; Madagascar; Seychelle Is¬
lands; Maidive Islands; Lakshadweep; Sri Lanka;
Andaman Islands; Indonesia; Philippines;
Ryukyu Islands; Caroline and Marshall Islands.

Periclimenes toloensis Bruce
(Figs 12d, f)

Periclimenes toloensisBmce, 1969:273-275;
1982: 258-264, figs 15-18; 1988b: 229-239.

Material examined. 1 spm., stn CP/17,
Wanaray Point, Trepang Bay, LWS, 15 October
1981, coll. A.J. Bruce, P. Horner, NTM
Cr.008733.9 spms, stn CP/26, Sandy Island No.
2, 7 m, 20 October 1981, P.N. Alderslade,
J.N.A. Hooper, NTM Cr.008590. Id", stn CP/27,
Sandy Island No. 2,10 m, 21 October 1981, coll.

J.N.A. Hooper et al., on unidentified hydroid,
NTM Cr.000159. 5 spms, stn CP/28, Sandy
Island No. 2,6-7 m, 22 October 1981, coll. J.R.
Hanley, on unidentified algae, NTM Cr.003186.
9 spms, same data as previous, on unidentified
hydroid, NTM Cr.003187. 3 spms, stn CP/37,
Table Head, 1-3 m, 3 May 1982, coll. H.K.
Larson, on whip coral Junceella sp., NTM
Cr.002937.2 spms, same data as previous, NTM
Cr.002941. 2 spms, same data as previous, on
soft coral Stereonephthya sp., NTM Cr.002957.

6 spms, stn CP/38, Table Head, 2-4 m, 4 May

1982, coll H.K. Larson et al., on soft coral
Sinularia polydactyla, NTM Cr. 001971.6 spms,
same data as previous, on soft coral
Neospongodes sp., NTM Cr.001973. 5 spms,
same data as previous, on Neospongodes sp.,
NTM Cr.001974. lei", I 9 , stn NY/4, McCluer
Island, 7 m. 16 October 1982, coll. A.J. Bruce,
NTM Cr.009275. 46 spms (7 ovig. 9 ), stn NY/ 8 ,
Oxley Island, LWS, 19 October 1982, coll. J.
Robinson, A.J. Bruce, NTM Cr.007771.12 spms
(2 ovig. 9 ), stn NY/9, Oxley Island, LWS, 20
October 1982, coll. A.J. Bruce, NTM Cr.009289.

7 spms (3 ovig. 9 ), stn CP/44, Table Head, 4-5 m,

11 May 1983, coll. N.L. Bruce, D. Staples, on
whip coral Junceella sp., NTM Cr.009349. I 9 ,
same data as previous, bopyridized, NTM
Cr.009354. 4 spms (1 ovig. 9 ), stn cp/46 ’ off
Table Head, 5 m, 12 May 1983, coll. N.L. Bruce,
NTM Cr.009340. lo", same data as previous,
NTM Cr.009341. 1 ovig. 9 , same data as previ¬
ous, NTM Cr.009342. 30 spms (17 ovig. 9 ),
same data as previous, NTM Cr.009351. 18
spms, same data as previous, NTM Cr.009352.

14 spms (1 ovig. 9 ), same data as previous, NTM
Cr.009355. 6 spms (1 spm. bopyridized), same
data as previous, NTM Cr.009357. 16 spms,
same data as previous, NTM Cr.009358. I 9 . 1
ovig. 9 , stn CP/ 68 , Coral Bay, 2-3 m, 19 May

1983. coll. N.L. Bruce, J.R. Hanley, NTM
Cr.009284. 110 spms (43 ovig. 9 ), stn CP/73,
Table Head, 6-8 m, 14September 1985, coll. C.
Hood, on hydroid Lytocarpus philippinensis, 2
spms with hemiarthrinid bopyrids (Cr.004218),
NTM Cr.004217. 1 spm., stn CP/78, Orontes
Reef, 19 m, 16September 1985,coll. R. Williams,
on soft coral Dendronephthya sp., NTM

Cr.007553. 9 ovig. 9 , stn CPV/ 8 , West of Barrow
Bay,LWS, 18 September 1985, coll J.R. Hanley,
M. Burke, on unidentified soft coral, NTM
Cr.007651.12 spms, stn CP/85. Orontes Reef, 10-

15 m, 19 September 1985, coll. L. Vail, C. Hood,
R. Williams, on coral Duncanopsammia sp., NTM

135

A.J. Bruce and K.E. Coombes

Cr.007612. 54 spms, stn CP/ 86 , Table Head, 8
m, 7 August 1986, coll. C. Johnson et al., on
unidentified gorgonian, NTM Cr.004090. 8
spms, same data as previous, on Lytocarpus
philippinensis , NTM Cr.004091. 39 spms (14
ovig. 9 , one bopyridized), same data as previous,
on Lytocarpus philippinensis, NTM Cr.004092.
24 spms (3 ovig. 9 ), same data as previous, on
Lytocarpus philippinensis , NTM Cr.004180.

Remarks. The specimens agree well with the
original description of Hong Kong material.
The rostrum is slightly deeper, with the ventral
border slightly more convex. The coxa of the
first pereiopod bears a large setose ventromedial
lobe and a similar smaller lobe is present on the
basis. The fourth thoracic stemite has a large
transverse triangular median tooth posteriorly.
The third sternite has a smaller similar lobe and
the fifth a transverse ridge with small submedian
teeth.

Australian distribution. Northern Territory:
East Point, Darwin (Bruce 1988b). Queensland:
Wistari Reef, Capricorn Islands.

Further distribution. Type locality: Ap Chau,
Hong Kong. Also known only from Zanzibar
and the Philippines.

Periclimenes venustus Bruce

Periclimenes venustus Bruce, 1990a: 230-
240, figs 1-6, 7a, 8 a; 1990b: 12.

Material examined. I 9 , stn CP/44, Table
Head, 4-5 m, 11 May 1983, coll. N.L. Bruce, D.
Staples, NTM Cr.009344. 1 ovig. 9 , paratype,
stn CP/64, Coral Bay, 3-4 m, 18 May 1983, coll.
N.L. Bruce, on coral Heliofungia actiniformis,
NTM Cr.000447. 2cf, I 9 , stn CP/71, Coral Bay,
2-5 m, 13 September 1985, coll. J.E.N. Veron,
on Heliofungia actiniformis, NTM Cr.003224.
la", 1 ovig. 9 , stn CP/72, Table Head, 3 m, 13
September 1985, coll. C. Hood, on unidentified
anemone, NTM Cr.003228.3 spms, stn CP/74,
Berkeley Bay, 3 m, 14 September 1985, coll. C.
Hood, J.E.N. Veron, on unidentified anemone,
NTM Cr.003229. 6 spms (1 ovig. 9 ), same data
as previous, on coral Heliofungia sp., NTM
Cr.003230. Id", 1 ovig. 9 , 1 juv., stn CP/76, Coral
Bay, 6 m, 15 September 1985. coll L. Vail, R.
Williams, J.E.N. Veron, NTM Cr.006340. I 9 ,
stn CP/81, off Turtle Point, Kennedy Bay, 2-5 m,
17 September 1985, coll. C. Hood, on unidenti¬
fied anemone, NTM Cr.006341. 29, stn CP/82,
Coral Bay,5-6 m, 18 September 1985,coll. P.N.
Alderslade, R. Williams, on Heliofitngia actini¬
formis, NTM Cr.006342. I 9 ,1 ovig. 9 , same data

as previous, NTM Cr.006345.1 spm., same data
as previous, NTM Cr.006353. 1 spm., on uni¬
dentified anemone, NTM Cr.006354. 1 spm.,
same data as previous, on Heliofungia actini¬
formis, NTM Cr.006355. 4 spms paratypes,
same data as previous, NTM Cr.006356.5 spms
paratypes, same data as previous, on unidenti¬
fied anemone, NTM Cr.006357. 1 ovig .9
Holotype, stn CP/97, Coral Bay, 12 August
1986,coll. C. Johnson, on unidentified anemone,
NTM Cr.004169. 1 spm., stn CP/98, Coral Bay,
13 August 1986, coll. C. Johnson, P. Davie,
NTM Cr.004170 (specimen destroyed).

Australian distribution. Type locality: Coral
Bay, Port Essington, Northern Territory (Bruce
1990a). Western Australia: Scott Reef; Abrolhos
Islands.

Further distribution. So far reported only
from the Philippines and Japan.

Periclimenes zanzibaricus Bruce

Periclimenes zanzibaricus Bruce, 1969: 62-
72, figs 26-29.

Material examined. 6 spms (2 ovig. 9 ), stn
CP/64, WalfordPoint, Coral Bay, 3-4m, 18 May
1983, coll. N.L. Bruce, on sea urchin Diadema
setosum, NTM Cr.000444.

Australian distribution. Northern Territory:
not previously recorded. Western Australia:
Geraldton.

Further distribution. Type locality: Mtoni,
Zanzibar. Also known from Kenya and the
Seychelle Islands.

Philarius gerlachei (Nobili)

Harpilius Gerlachei Nobili, 1905: 160; 1906:
45, pi. 4, fig. 10.

Philarius gerlachei - Holthuis 1952: 152-
153, fig. 69.

Material examined. 1 juv., stn CP/33, Port
Bremer, 6 m, 1 May 1982, coll. J.N.A. Hooper,
P.N. Alderslade, NTM Cr.009318. 1 ovig. 9 , stn
NY/4, McCluer Island, 7 m, 16 October 1982,
coll. A.J. Bruce, NTM Cr.009278. lei", stn CP/
82, Coral Bay, 5-6 m, 18 September 1985, coll.
P.N. Alderslade, R. Williams, on coral Acropora
sp., NTM Cr.007579. lcf, 1 ovig. 9 , same data as
previous, on Acropora digitifera, NTM
Cr.007609. 1 ovig. 9 , same data as previous, on
Acropora sp., NTM Cr.010549. la", I 9 , stn CP/
88 , Orontes Reef, 8-10 m, 9 August 1986, coll.
R.C. Willan, C. Johnson, on Acropora sp., NTM
Cr.004101.

136

The palaemonoid shrimp fauna of the Cobourg Peninsula

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island; One Tree Island; Diamond Islet, Willis
Island; Restoration Rock.

Further distribution. Type locality: Arzana
Island, Persian Gulf. Also known from the Red
Sea; Oman; Kenya; Zanzibar; Tanganyika;
Mozambique Channel; Comoro Islands;
Seychelle Islands; La Reunion; Gulf of Manaar;
Indonesia; Philippines; Ryukyu Islands; Marshall
Islands; Solomon Islands; Fijian Islands; Kiribati
and Samoan Islands.

Philarius imperialis (Kubo)

Harpilius imperialis Kubo, 1940b: 1-4, figs
1-3.

Philarius imperialis - Holthuis 1952: 125; -
Bruce 1983a: 45.

Material examined. Id", I 9 , stn CP/20,
Walford Point, Coral Bay, LWS, 17 October
1981, coll. A.J. Bruce et al ., NTM Cr.000132.2
spms, stn CPI 40, Orontes Reef, 3 m, 5 May 1982,
coll. A.J. Bruce, on coral Acropora sp., NTM
Cr.000281. 1 spm., same data as previous, coll.
H.K. Larson, NTM Cr.001237. lcf, 1 ovig. 9 ,stn
CP/61,Coral Bay,4m, 17May 1983,coll.N.L.
Bruce, J.R. Hanley, on Acropora sp., NTM
Cr.010516. 2 spms, stn CP/64, Walford Point,
Coral Bay, 3-4 m, 18 May 1983, coll. N.L. Bruce,
on Acropora sp., NTM Cr.010245.1 ovig. 9 , stn
CP/81, offTurtle Point, Kennedy Bay, 2-5 m, 17
September 1985, coll. C. Hood, on Acropora
millepora, NTM Cr.007581. Id", stn CP/95,
Coral Bay,5-6m, 12 August 1986, coll. P. Davie
et al., on Acropora sp., NTM Cr.004153.

Remarks. The specimens NTM Cr.000132
and NTM Cr.010516 were found in association
with Periclimenes lutescens specimens.

Australian distribution. Northern Territory:
Coral Bay, Port Essington (Bruce 1983a).
Queensland: Heron Island, Restoration Rock.

Further distribution. Type locality: Bonin Is¬
land. Also recorded from the Red Sea; Kenya;
Zanzibar, Tanganyika; La Reunion; Singapore;
Indonesia; Caroline Islands and Marshall Islands.

Typton dentatus Fujino and Miyake

Typton dentatus Fujino and Miyake, 1969b:
80-84, figs 1-2.

Material examined. Id", I 9 , stn CP/64,
Walford Point, Coral Bay,3-4m, 18May 1983,
coll. N.L. Bruce, NTM Cr.009286.

Remarks. The specimens were collected from
smashed coral rubble, presumably from a sponge
host.

Australian distribution. Northern Territory:
not previously recorded. Queensland: Heron
Island.

Further distribution. Type locality: Yoron-
jima, Ryukyu Islands. No other records.

Anchistioididae Gurney, 1938

Anchistioides willeyi (Borradaile)

(Figs 12g, h)

Palaemonopsis willeyi Borradaile, 1899:410,
pis 36-37, fig. 7.

Amphipalaemon willeyi - Nobili 1901: 5.

Anchistioides willeyi - Gordon 1935: 345,
figs 23a, 24a; - Bruce 1978b: 286-287, fig. 44;
1988b: 227; 1991: 269-272, figs 3g, 29-30.

Material examined. 2 juv., stn HL82-60,
New Year Island, surface, night-light, 13 Octo¬
ber 1982, coll. H.K. Larson, NTM Cr.000322.
Id", stn HL82-73, Oxley Island, surface, night-
light, over 9-10 m, 18 October 1982, coll. H.K.
Larson, NTM Cr.000324.

Remarks. The capture of these specimens at
surface night-lights represents the first occur¬
rence of this phenomenon in the Indo-West
Pacific species of this genus. The new moon
occurred on 17 October 1982, so that specimens
NTM Cr.000322 were collected three days be¬
fore the new moon and NTM Cr.000324, one
day after. This occurrence, in the case of
Anchistioides antiguensis (Schmitt) in the Car¬
ibbean region, has been documented in detail by
Wheeler and Brown (1936). Swarming was re¬
ported to occur shortly after sunset at the time of
the new moon. At that time the normal associa¬
tion of Anchistioides species with sponge hosts
was not known (there still appear to be no
confirmed reports of A. antiguensis being found
in sponges). It is interesting to note that in most
of the Bermudan specimens, the females were
ovigerous, so that the swarming was not a pre¬
mating phenomenon. Some of the females cap¬
tured clearly had ova in the point of eclosion. It
has also been reported by Wheeler (1944) that
the stomach contents of A. antiguensis may
contain setae of the polychaete worm Perinereis
melanocephala M’lntosh and that captive speci¬
mens of the shrimps feed readily on the epitokes
of Perinereis. How the swarming shrimps re¬
gain their sponge hosts is a mystery, but their
glassy transparency and almost complete lack of

137

A.J. Bruce and K.E. Coombes

chromatophores may help to expedite their re¬
turn.

Two specimens (NTM Cr.000322) appear
juvenile, with CL 3.9 and 4.8 mm, and have
chelae distinctly of the long, slender-fingered
type (Bruce 1978b, 1991). The distal teeth,
medial to the sharp cutting edge are highly
acute, the proximal denticles are rounded. The
larger male (NTM Cr.000324),CL 8.1 mm, may
be adult and has similar chelae. In both lots, each
finger has a pair of very long rigid simple setae,
that become erect on the opened fingers, but
depressed when the fingers are closed. These
appear to be autapomorphic characters, lacking
in all other palaemonoid genera. These setae

may be correlated with the pelagic feeding hab¬
its mentioned above. These features, and the
unusual morphology of the mandibles, with the
unusual proximal segments of the third
maxilliped (Bruce 1991) and particularly the
unusual larval morphology (Gurney 1936) sug¬
gest that the Anchistioididae are not closely
related to the rest of the Palaemonoidea.

Australian distribution. Northern Territory:
Darwin Harbour (Bruce 1988b). Also known
from Queensland, Heron and One Tree Islands,
and from Moreton Bay (Bruce 1983c).

Further distribution. Type locality: Ralun,
New Britain. Also recorded from Zanzibar; Tan¬
ganyika; Kenya; Madagascar; Maidive Islands;

TABLE 3. Comparison of Cobourg Peninsula and Singapore pontoniine faunae. Species recorded only from the Cobourg
Peninsula or Singapore are indicated by •.

Taxon Cobourg

Peninsula

Singapore

Host

Palaemoninae Exopalaemon stylifera (H. Milne Edwards 1840)

—

Leander tenuicornis (Say, 1818)

Leandrites celebensis (De Man, 1811)

Leandrites deschampsii (Nobili, 1903)

Leandrites stenopus Holthuis, 1950

Leptocarpus potamiscus (Kemp, 1917)

Palaemon concinnus Dana, 1852

Palaemon debilis ( Dana, 1852)

Palaemon semmelinckii (De Man, 1881)

Palaemon serrifer (Stimpson, 1860)

Urocaridella antonbruunii (Bruce, 1967)

(fish - cleaner)

Urocaridella urocaridella (Holthuis, 1950)

Urocaridella sp.

(fish - cleaner)

Pontoniinae Anapontonia denticauda Bruce, 1967

Scleractinia: Ocutinidae

Anchistus australis Bruce, 1977

Bivalve mollusca

Anchistus custos (ForsskSl, 1775)

Bivalve mollusca: Pinnidae

Anchistus miersi (De Man, 1888)

Bivalve mollusca: Tridacnidae

Chernocaris placunae Johnson, 1967

Bivalve mollusca: Placunidae

Conchodytes monodactylus Holthuis, 1952

Bivalve mollusca: Pinnidae

Coralliocaris graminea (Dana, 1852)

Scleractinia: Acroporidae

Coralliocaris viridis Bruce, 1974

Scleractinia: Acroporidae

Dasella ansoni Bruce, 1983

•

Tunicata

Hamodattylus boschmai Holthuis, 1952

Hamodactylus noumeae Bruce, 1970

Hamopontonia corallicola Bruce, 1970

Scleractinia: Actinaria

Hamopontonia essingloni Bruce, 1987

•

Scleractinia: Pocilloporidae

Harpiliopsis beaupresii (Audouin, 1852)

Scleractinia: Pocilloporidae

Ischnopontonia lophos (Barnard. 1962)

Scleractinia: Oculinidae

Onycocaris quadratophthalma (Balss, 1921)

Porifera

Palaemonella pottsi (Borradaile, 1915)

Crinoidea

Palaemonella rolumana (Borradaile, 1898)

Palaemonella spinulata Yokoya, 1935

Periclimenaeus arabicus (Caiman, 1939)

Porifera

Periclimenaeus orontes Bmce. 1989

•

Porifera

Periclimenaeus serrula sp. nov.

•

Porifera

Periclimenaeus solitus sp. nov.

•

Porifera

Periclimenaeus stylirostris Bruce, 1969

Porifera

Periclimenaeus Iridentatus (Micrs, 1844)

Tunicata

Periclimenella spinifera (De Man, 1902)

Periclimenes affinis (Zehntner, 1894)

Crinoidea

Periclimenes akiensis Kubo, 1936

138

The palaemonoid shrimp fauna of the Cobourg Peninsula

Singapore; BomeoBank; Indonesia; South China
Sea; Philippines, and New Caledonia.

DISCUSSION

The collection of over 1500 specimens of
palaemonoid shrimp from the Cobourg Penin¬
sula region provides some interesting data on
the relative abundance of some of the commensal
species, although collections were made only on
a quanlitative basis. The commensal species
collected greatly outnumber the ‘free-living’
species, and illustrate the density with which
some species can occur. Thus, the most abun¬
dant ‘free-living’ species was the ubiquitous

Palaemonella rotumana (52 spms), which has
often been noted as one of the most common and
most widely distributed pontoniine species, oc¬
curring throughout most of the Indo-West Pa¬
cific region and now also occurring in the east¬
ern Mediterranean Sea, while also extending to
a depth of 126-128m (Bruce 1970c). These
specimens were almost invariably collected in
lots of 1-2 specimens only. Specimens of this
species were caught, in many cases almost acci¬
dentally, in the search for other species, as well
as from rotenone stations. The total numbers of
the commensal species naturally depend upon
the number of potential hosts examined. Num¬
bers per host vary considerably, with one male

TABLE 3 (cont.). Comparison of Cobourg Peninsula and Singapore pontoniine faunae. Species recorded only from the
Cobourg Peninsula or Singapore are indicated by •.

Taxon Cobourg Singapore Host

Peninsula

Pontoniinae

Periclimenes alegrias Bruce, 1987

•

Crinoidea

(cont.)

Peridimenes amymone De Man, 1902

Scleractinia

Periclimenes anacamhus Bruce, 1989

Periclimenes brevicarpalis (Schenkel, 1902)

Actinaria

Periclimenes cobourgi sp. nov.

•

Scleractinia

Periclimenes commensalis Borradaile, 1915

Crinoidea

Periclimenes cristimanus Bruce, 1965

Echinoidea

Periclimenes digitalis Kemp, 1922

Periclimenes diversipes Kemp, 1922

Scleractinia

Periclimenes elegans (Paulson, 1875)

Periclimenes galene (Holthuis, 1952)

Hydroida

Periclimenes grandis (Stimpson, 1860)

Periclimenes holthiusi Bruce, 1969

Actinaria: Scleractinia

Periclimenes incertus Borradaile, 1915

Periclimenes indie us (Kemp, 1915)

Periclimenes investigatoris Kemp, 1922

Periclimenes johnsoni Bruce, 1987

•

Periclimenes kempi Bruce, 1969

Alcyonacea

Periclimenes lanipes Kemp, 1922

Ophiuroidea

Periclimenes lutescens (Dana, 1852)

Scleractinia

Periclimenes madreporae (Bruce, 1969)

Periclimenes magnificus Bruce, 1979

Scleractinia

Periclimenes mahei Bruce, 1969

Scleractinia

Periclimenes nilandensis Borradaile, 1915

Gorgonacea

Periclimenes novaecaledoniae Bruce, 1968

Crinoidea

Periclimenes obscurus Kemp, 1922

Periclimenes parvus Borradaile, 1898

Periclimenes psamatlie (De Man, 1902)

Gorgonacea

Periclimenes seychellensis Borradai le, 1915

Periclimenes sinensis Bruce, 1969

Periclimenes soror Nobili, 1904

Asteroidea

Periclimenes tenuipes Borradaile, 1898

Periclimenes toloensis Bruce, 1969

Hydroida

Periclimenes venustus Bruce, 1990

Scleractinia

Periclimenes zanzibaricus Bruce, 1969

Echinoidea

Philarius gerlachei (Nobili, 1905)

Scleractinia

Philarius imperialis (Kubo, 1940)

Scleractinia

Platycaris latirostris Holthuis, 1952

Scleractinia: Oculinidae

Typton dentatus Fujino & Miyake, 1969

Porifera

Anchistioididae

Anchistioides willeyi (Borradaile, 1899)

Porifera

Hymenoceridae

Phyllognathia ceratophthalma (Balss, 1913)

139

A.J. Bruce and K.E. Coombes

and one female being the normal occupancy of
bivalve mollusc hosts. This is also normal in
tunicate hosts and contrasts strongly with the
numbers of specimens found on some coelenterate
or sponge hosts. Thus, although most host speci¬
mens usually only provided a small number of
commensal shrimps, others contained large num¬
bers, such as the 110 specimens of Periclimenes
toloensis on a single hydroid host, Lytocarpus
philippinensis. Similarly, one crinoid host pro¬
vided 64 associated specimens of Periclimenes
affinis.

Faunistic comparisons with other tropical ar¬
eas present problems as the protocols of collec¬
tions differ markedly. Bruce (1981) studied the
fauna of Heron Island, Queensland, and re¬
corded well over 100 species of pontoniine
shrimp. This study illustrates the results of
prolonged study of a small coral reef, lacking
many of the major habitats found elsewhere in
the tropics. At the other extreme, the station
positions of the Siboga and Snellius expeditions
are diffusely spread over much of the Indonesian
archipelago, extending into deep-sea waters,
and so are not comparable. The best available
data for comparison with the Cobourg Peninsula
fauna is that provided for Singapore by Johnson
(1961, 1976), the results of detailed study over
many years of a relatively restricted area, with a
wide variety of habitats. A comparison of the
faunas of the two localities is presented in the
following table.

The two faunas combined, present a faunistic
total of 82 palaemonoid species, of which the
Cobourg Peninsula has 61 species (74%) and
Singapore 45 (55%), with only 24 (29%) occur¬
ring in both regions. Further collecting would
surely increase all these numbers, as several
common Indo-West Pacific species, such as
Jocaste lucina and Jocaste japonica and further
Coralliocaris spp (such as Coralliocaris
superba), all common in Acropora corals, and
Harpiliopsis depressa abundant in pocilloporid
corals, have not yet been recorded from either
region. Although there is a large element of
chance in collecting shrimps, the results of
surveys will reflect the methods employed. The
Singapore fauna contains a number of species
probably from trawl-caught samples, e.g.
Urocaridella urocaridella, Periclimenes lanipes,
while the Cobourg Peninsula fauna probably
lacks some Anchistus species, such as A.demani,
A. miersi, because theirhosts, Tridacna spp, are
protected and were not sampled. It should also

be noted that sponges, bivalve molluscs and
tunicates, all potential shrimp hosts, were also
inadequately sampled. Bruce (1981) reported
that a well developed coral reef may have over
100 species of pontoniine shrimps alone, al¬
though palaemonine shrimps are usually very
poorly represented in this biotope. Bruce (1990b)
reported the presence of 168 species of pontoniine
shrimps in the total Australian fauna. The pres¬
ence of 55 pontoniine species in the Gurig
Marine Park, which accounts for 32% of the
Australia fauna, reflects the rich marine fauna
present in this area, and is interesting in view of
the relatively poor development of its coral reefs.
The central waters of Port Essington were not
adequately surveyed by trawl or dredge, meth¬
ods that would certainly have increased the
species diversity. The low representation of
palaemonine shrimps at Port Essington may be
attributed to the lack of any major permanent
outflows of freshwater, such as occur in the
Singapore region. However, Leandertenuicornis
and Leandrites celebensis do both occur in
Northern Territory waters, and so could be found
in the Port Essington region in due course.

A short-term study of the caridean fauna of
Cartier-Hibemia reefs, off north western Aus¬
tralia, (Bruce 1992) provides some comparative
data on the coral reef fauna. The collection
included 45 palaemonoid species (Palaemoninae,
2; Pontoniinae, 42; Gnathophyllidae, 1). Of
these, only 12 (15%) occur in common with the
Cobourg Peninsula material, a marked contrast
with the degree of overlap between the Cobourg
and the Singapore fauna. The differences be¬
tween the Cobourg and Cartier-Hibemia reef
faunas are attributed to the relatively greater
amount of collecting carried out in the Cobourg
Peninsula and the greater degree of coral reef
development at Cartier-Hibemia reefs.

ACKNOWLEDGEMENTS

We are most grateful to the Heritage Com¬
mission for their financial support of the Port
Essington survey. The help of the master and
crew of the FV Alegrias are also much appreci¬
ated, together with the field assistance of nu¬
merous colleagues who participated in the study.
Dr Penny Berents kindly provided the speci¬
mens of Periclimenaeus and Allopontonia iaini
from Elizabeth and Middleton Reefs for re¬
examination. The Conservation Commission of

140

The palaemonoid shrimp fauna of the Cobourg Peninsula

the Northern Territory provided the necessary
permits to carry out the survey. The assistance
of Loma Gravener in the preparation of this
report is also greatly appreciated.

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A.J. Bruce and K.E. Coombes

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Accepted 1 June 1995

144

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:145-150

TWO NEW SPECIES OF THE GENUS ANISOMYSIS (CRUSTACEA:
MYSIDACEA) FROM NORTHERN AUSTRALIA.

MASAAKI MURANO

Department of Aquatic Biosciences, Tokyo University of Fisheries,

4-5-7 Konan, Minato-ku, Tokyo 108, Japan.

ABSTRACT

Nine species and subspecies of Anisomysis have been recorded from Australian waters
to date. These are A. bipartoculata, A. gracilis, A hispida, A.incisa, A, laticauda, A.
mixta australis, A. mullini, A. pelewensis and A. robustispina. The present two new
species, therefore, are the tenth and eleventh species of Anisomysis, and their occur¬
rence is the first record from Northern Territory waters. Anisomysis spatulispina n. sp.
is distinguished from other species of the genus by the shape and armature of the telson.
Anisomysis nana n. sp. is distinguished from allied species by having a rounded
rostrum and the telson with a deep, U-shaped cleft.

Keywords: Crustacea, Mysidacea, Anisomysis, Anisomysis spatulispina n. sp.,
Anisomysis nana n. sp. Northern Territory, Australia.

INTRODUCTION

Two new species of the genus Anisomysis
Hansen describedherein are based on specimens
which were collected in the Northern Territory of
Australia and stored in the Northern Territory
Museum (NTM). From Australia, nine species of
Anisomysis have been recorded as follows: A.
mixta australis Zimmer from Victoria (Zimmer
1918); A. mixta australis from New South Wales
(Tattersall 1927); A. mixta australis from South
Australia (Tattersall 1927); A. hispida Pillai (Pil-
lai 1973), A. bipartoculata\\,A.gracilis?anain-
punnayil andA. robustispinaPanampuimayil from
Western Australia (Panampunnayil 1984); A. in-
cisa Tattersall, A. laticauda Hansen (Tattersall
1936), A. mixta australis, A. pelewensis Ii (Fen¬
ton 1986) and A. mullini Murano from Queens¬
land (Murano 1987). The present report is the
first record of Anisomysis from Northern Territo¬
ry waters.

SYSTEMATICS

Anisomysis Hansen, 1910
Anisomysis spatulispina n. sp.

(Fig. 1A-I)

Type Material. HOLOTYPE - adult male,
NTM Cr.004319, 5.9 mm, Coral Bay, Port

Essington, Northern Territory, Australia,
11°11.3’S 132°11.5’E,4m, 12September 1985,
collected by A.J. Bruce.

Other material. 1 immature male, NTM
Cr.004130,3.4 mm, Coral Bay, Port Essington,
7 m, 11 August 1986.

Description. Rostrum triangular, with nar¬
rowly rounded apex overreaching base of anten-
nular peduncle, lateral margin of rostrum con¬
vex (Fig. 1A); anterolateral comer of rostrum
rounded; posterior margin of carapace emarginate
dorsally, leaving last thoracic somite uncovered.

Eye large; cornea globular, wider than eyestalk;
eyestalk with prominent groove at junction of
cornea, without any papilliform process on dor¬
sal surface (Fig. 1 A).

Antennular peduncle robust, first segment as
long as third, armed on distolateral comer with
several setae, one longerand thickerthan others,
third segment with large processus masculinus
(Fig. 1A).

Antennal scale extending beyond distal end of
antennularpeduncle butnottoapex ofprocessus
masculinus, slender, 7 times longer than broad,
outer margin very slightly concave, inner mar¬
gin convex, setose all round, distal suture well
marked (Fig. IB). Antennal peduncle short, less
than half as long as scale (Fig. IB).

First segment of mandibular palp short, with
triangular process on inner margin; second seg-

145

M. Murano

Fig. 1. Anisomysis spatulispina n. sp.: A, anterior end; B, antenna; C, mandibular palp; D, endopod of fourth thoracic
limb; E, endopod of Fifth thoracic limb; F, fourth pleopod; G, uropod and telson; H, right distal lobe of telson;
I, telson of immature male.

146

Two new species of the genus Anisomysis

ment with four prominent flagellate denticles
along inner margin, proximal two largest, outer
margin very slightly concave, with 23 barbed
setae arranged at rather regular intervals; third
segment subrectangular, inner margin with five
barbed setae, distal margin with seven setae of
three kinds, outer margin with single short and
simple seta at distal end (Fig. 1C).

Endopods of third to eighth thoracic limbs of
usual form in this genus, carpopropodus undi¬
vided into subsegments (Fig. ID, E). Flagella of
thoracic exopods segmented to 7-8 .

Abdomen with first four somites subequal;
fifth somite 1.3 times longer than the fourth;
sixth somite 1.4 times longer than the fifth.

Fourth male pleopod biramous; endopod re¬
duced to small lobe with several setae; exopod
extremelyelongated, reachingdistalendoftelson,
3-segmented, first segment3.0 times longerthan
second segment, third segment 1.2 times longer
than second segment, with two terminal setae of
same length (Fig. II 7 ).

Uropod slender, setose all round; endopod
straight, twice as long as telson; exopod slightly
curved outward, overreaching endopod by about
1/5 of its length (Fig. 1G).

Telson short, 3/4 length of sixth abdominal
somite, 1.6 times longer than broad at base,
constricted at distal third, narrowest part about
half as wide as base, distal part posterior to
constriction again becoming broader and bilobed;
distal sinus deep, 1/4 of telson length; lateral
margin of telson naked in proximal 2/5, armed in
distal 3/5 with about 10 spines, four or five being
located before constriction; apices and sinus
with 18 spines, flattened and rounded at apex
(Fig. 1G, H).

Remarks. Anisomysis spatulispina n. sp. re¬
sembles Anisomysis hosakaiMurano, 1990, and
Anisomysis gracilis Panampunnayil, 1984, in
having the mandibular palp with flagellate
denticles on the inner margin of second segment
and in the telson with both lateral constriction
and distal sinus. From A. hosakai, however, the
new species differs in telson shape, i.e. the telson
is clearly constricted in the new species, whereas
in A. hosakai the constriction is so slight that it is
barely recognizable, and the distal sinus is deeper
(1/4 of telson length) than in A. hosakai (1/5 of
telson length). From A. gracilis, the new species
is also distinguishable in the following: (1)
antennal scale 7.0 times as long as broad and not
extending beyond apex of processus masculinus

in the new species, while 10 times as long as
broad and far extending beyond processus
masculinus in A. gracilis; (2) second segment of
mandibular palp with four flagellate denticles in
the new species versus seven in A. gracilis ; (3)
exopod of fourth male pleopod with second
segment shorter than the third in the new species,
while longer in A. gracilis; (4) constriction of
telson half as broad as base in the new species,
while it is 1/4 as broad in A. gracilis; and (5)
distal sinus of telson 1/4 as long as telson in the
new species, while it is 1/13 as long in A. gracilis.

Etymology. The specific name is derived from
the shape of marginal spines on the telson sinus.

Anisomysis nana n. sp.

(Fig. 2 A-F)

Type material. HOLOTYPE - adult female
with fully developed marsupium, NTM
Cr.005918,3 m, Nightcl iff Beach, Darwi n. North -
em Territory, Australia, reef flat pool, 11 Sep¬
tember 1987, collected by D. Sachs.

Description. Carapace with anterior margin
broadly rounded, barely extending to base of
antennular peduncle, partly covering eyestalk
(Fig. 2A); anterolateral comer rounded; poste¬
rior margin emarginate, leaving last thoracic
somite exposed dorsally.

Eye large with globular cornea, extending
laterally beyond lateral margin of carapace (Fig.
2A).

Antennular peduncle rather slender, first seg¬
ment as long as third segment (Fig. 2 A). Antennal
scale reaching distal end of antennular peduncle,
slender, nearly straight, about 7.0 times longer
than broad, both outer and inner margins setose,
distal suture absent. Antennal peduncle short,
not extending to middle of scale, second and
third segments wider than long (Fig. 2B).

Mandibular palp foliated, second segment 3.0

times as long as broad, without flagellate denticles

on inner margin, third segment more than 3.0
times as long as broad, subrectangular.

Endopod of seventh thoracic leg with ischium
shorter than merus, merus longer than
carpopropodus and dactylus combined,
carpopropodus not divided into subsegments,
dactylus about 1/4 of carpopropodus in length,
terminating in slender nail; exopod distal to
basal plate 8-segmented (Fig. 2D). Eighth tho¬
racic limb considerably smaller than seventh,
merus of endopod shorter than carpopropodus

147

M. Murano

Fig. 2. Anisomysis nana n. sp. : A, anterior end; B, antenna; C, mandible; D, seventh thoracic limb; E, eighth
thoracic limb; F, uropod and telson.

148

Two new species of the genus Anisomysis

and dactylus combined, without setae on both
inner and outer margins except two setae at outer
distal comer; exopod distal to basal plate 7-
segmented (Fig. 2E).

Sixth abdominal somite 1.5 times longer than
the fifth, 1.5 times longer than broad.

Uropod setose all round; exopod extending
posteriorly beyond telson for 1/4 of its length,
slightly curved outward; endopod tapering, as
long as exopod, without spines in statocyst re¬
gion (Fig. 2F).

Telson slightly longer than last abdominal
somite, 2.5 times as long as broad at base, very
deeply cleft for about half its length, cleft very
wide, U-shaped and unarmed, distal part of telson
forming two-pronged fork; lateral margin armed
along distal 2/3 with 9-11 similar-sized spines,
six or seven of which are on fork, each fork apex
with single spine considerably longer than lat¬
eral spines.

Remarks. The new species is characterized
by the two-pronged telson, which has a deep
cleft without spines and each apex terminating in
a single spine. Such a telson is known in five
species in the genus Anisomysis: A. bifurcata
Tattersall, 1912, A. megalops (Illig, 1913), A.
pelewensis li, 1964, A. minuta Liu and Wang,
1983, and A..Panampunnayil, 1993. The
present new species is most closely related to A.
spinata, from the Lakshadweep Archipelago in
the Arabian Sea, in that the telson has a deep and
U-shaped cleft. The new species, however, is
distinguishable from the latter as follows: (1)
anterior margin of carapace broadly rounded,
while it is triangular with obtusely pointed apex
in A. spinata; and (2) lateral margin of telson
armed with 9-11 spines,6-7ofwhich arise on the
prong, whereas in A. spinata the number of
spines is 6, only 2 of which are on the prong.

The new species is also allied to A. megalops
in possessing a deep cleft in the telson, but differs
from the latter which ischaracterized by a pointed
rostrum, the eye having a small process on its
stalk, the antennal scale extending beyond the
distal end of the antennular peduncle, the telson
cleft being rather V-shaped and the telson with 2
or 3 spines on the lateral margin of prong (Illig,
1913, 1930). Anisomysis nana n. sp. is easily
distinguished from the remaining three species
by the shape and armature of the telson.

Etymology. The species name is derived from
the very small-sized body.

ACKNOWLEDGEMENT

I wish to express my sincere thanks to Dr. A.

J. Bruce, formerly of the Museum of Northern

Territory of Australia, for giving me the oppor¬
tunity of examining the present material.

REFERENCES

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Liu Ruiyu and Wang Shaowu. 1983. On three new
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waters of Akajima Island, Ryukyu Islands. Jour¬
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212 .

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longing to the genus Anisomysis (Crustacea,
Mysidacea) and a new record of Anisomysis
bipartoculata from the Indian Ocean. Journal of
Plankton Research 6: 943-952.

Panampunnayil, S. U. 1993. Two new species of
Anisomysis (Crustacea-Mysidacea) from
Lakshadweep archipelago. Journal of Plankton
Reseaarch 15: 1141-1148.

Pillai, N.K. 1973. Mysidacea of the Indian Ocean.
Handbook to the international zooplankton col¬
lections. Papers on the Zooplankton Collections
of the 1IOE 7: 1-125.

Tattersall, W. M. 1912. On the Mysidacea and
Euphausiacea collected in the Indian Ocean dur¬
ing 1905. The Percy Sladen Trust Expedition to
the Indian Ocean in 1905, vol. 4, No. 9. Transac¬
tions of Linnean Society of London, Zoology,
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Tattersall, W. M. 1927. Australian opossum shrimps.
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149

M. Murano

tific reports. London, British Museum (Natural
History) 5: 143-176.

Zimmer, C. 1918. Neue und wenig bekannte
Mysidaceen des Berliner Zoologischen Muse¬

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Accepted 18 July, 1995

150

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:151 -176

SYSTEMATICS AND BIOGEOGRAPHY OF THE GENUS IANTHELLA
(DEMOSPONGIAE: VERONGIDA: IANTHELLIDAE)

IN THE SOUTH-WEST PACIFIC.

PATRICIA R. BERGQUIST' AND MICHELLE KELLY-BORGES 1 - 2

'Zoology Department, School of Biological Sciences, University of Auckland,
Private Bag 92019, Auckland, New Zealand.

2 Present address: Department of Zoology Natural History Museum,

Cromwell Road, London, SW7 5BD, United Kingdom.

ABSTRACT

Three new species of lanthella Gray (Demospongiae; Verongida; Ianthellidae) from
Papua New Guinea and north-western Australia are described. lanthella flabelliformis
Gray and I. basta Gray are redescribed following reference to type material and to new
material. Diagnostic characters of lanthella species are stated and subjected to
phylogenetic analysis in order to evaluate the uniformity of the genus. The systematics
and biogeography of lanthella are reviewed.

Keywords: Porifera, Demospongiae, Verongida, Ianthellidae, lanthella, new species,
taxonomy.

INTRODUCTION

The verongid family Ianthellidae contains
sponges in which the fibre skeleton is most
commonly compressed into two dimensions, in
an arrangement which isalwaysareticulation of
primary fibres radiating from a constricted base
of attachment and supported by abundant con¬
necting fibres. The genera lanthella Gray and
Anomoianthella Bergquist differ in the degree
of regularity, fasciculation, and coarseness of
the fibre skeleton. Fibres consist of a diffuse pith
surrounded by a concentrically laminated bark,
into which spongocytes are incorporated. The
choanosomal construction of all Ianthellidae is
cavernous, but the region can be reinforced
locally with collagen. Choanocyte chambers are
eurypylous, sometimes markedly elongate.

The genus lanthella contains thin, frequently
large fan or vase-shaped sponges, which display
the typical verongid range of colours, yellow,
orange, green, blue and purple. In lanthella the
skeleton, which makes up the bulk of the sponge
body, is developed in two dimensions; it can be
a complex reticulation of fascicles, or an
anastomosing network of single fibres. Exten¬
sions of the fibres at right angles to the two

dimensional plane of the sponge are present in
several species.

Six species of lanthella have been described
by Lendenfeld (1889) and Polejaeff (1884). Of
these species, only /. flabelliformis (Pallas) and
I. basta (Pallas) were considered to be valid
(Bergquist 1980), although there is considerable
confusion in the past literature regarding the
diagnostic characters that differentiate these
easily recognised species, lanthella ianthella de
Laubenfels has been transferred to the genus
Verongula (Wiedenmeyer 1977), and lanthella
ardis de Laubenfels is a synonym of Aiolochroia
crassa (Bergquist 1980, 1995). Ianthella
concentrica Hyatt and 1. homei Gray have been
declared unrecognisable by Bergquist (1980). In
this communication we report three new species
of lanthella from northern Australia, eastern
Australia and Papua New Guinea, and redescribe
l. flabelliformis and I. basta.

METHODS

Colournotation forpreserved and living speci¬
mens follow Munsell (1942). Collections were
made by the authors unless otherwise stated. To

151

P.R. Bergquist and M. Kelly-Borges

examine skeletal structure, sections of sponge
tissue were macerated in 10 % sodium
hypochlorate for 2-5 minutes, washed thoroughly
in water and stored in 70 % ethanol. Skeletons
were mounted on aluminium stubs and sputter-
coated with 10 nm gold. Skeletons were exam¬
ined and photographed using a Phillips 505
scanning electron microscope at 20 kV. The
holotype of Ianthella reticulata sp. nov. has
been deposited in the Australian Museum (AM),
Sydney, Australia, and the registration numbers
are cited in the text. The holotypes of Ianthella
labyrinthus sp. nov. and /. quadrangulata sp.
nov. are in the Northern Territory Museum
(NTM), Darwin, Australia, and the Natural
History Museum (BMNH), London, United
Kingdom, respectively. Other abbreviations -
MNHN: Museum National d’Histoire Naturelle,
Paris, France; AIMS: Australian Institute of
Marine Sciences,Townsville, Australia; UAZA;
Zoology Department, University of Auckland,
Auckland, New Zealand; NSRC (UPNG): Natu¬
ral Science Resource Centre, University of Papua
New Guinea, Port Moresby, Papua New Guinea;
CRRF: Coral Reef Research Foundation, Micro¬
nesia; 0CDN: specimen sample numbers for
United States National Cancer Institute shal¬
low-water collection program contracted to
CRRF. A complete collection of all 0CDN speci¬
mens is located at the Smithsonian Institution
(United States National Museum); Q66C: speci¬
men sample numbers for United States National
Cancer Institute shallow-water collection pro¬
gram previously contracted to AIMS. This latter
collection is destined to be located at the Queens¬
land Museum, Brisbane, Australia.

SYSTEMATICS

Order Verongida Bergquist
Family Ianthellidae Hyatt
Genus Ianthella Gray

Ianthella Gray, 1869: 49; Hyatt, 1875: 407;
Polejaeff, 1884: 37;Lendenfeld, 1888:23,1889:
683; Wilson, 1925: 474; de Laubenfels, 1936:
31, 1948: 154, Bergquist, 1980: M3-503.
Basta Oken, 1815: 77; Burton, 1934: 596.
Haddonella Sollas, 1903: 557.

Type species. Spongia flabelliformis Pallas
1776: 380 (by subsequent designation, Topsent
1905).

Diagnosis. Ianthellidae in which the fibre
skeleton is compressed in a single plane to
produce a single, bilamellate or multilamellate
fan or vase, frequently of very large size. The
skeleton makes up the bulk of the sponge body
and can be a rectangular reticulation of fascicles
crosslinked by secondary Fibres, or a simple
reticulation of anastomosing fibres, developed
in two dimensions. Fibre outgrowths at right
angles to the basic two dimensional reticulum
are present in several species. Fibres are deep
reddish purple and typically consist of bark with
spongocytes in concentric annuli, which sur¬
round a pith that makes up between 20 to 50% of
the total fibre diameter. The bark immediately
surrounding the pith is compact and contains
the great bulk of the spongocytes. The outer
region of the fibre is less compact, and contains
fewer spongocytes. The choanosome is cavern¬
ous with large, sac-shaped eurypylous choanocyte
chambers. The mesohyl of the choanosome is
usually lightly to moderately reinforced with
collagen, but the ectosomal region has strong
collagen deposition. Sand and detritus can be
incorporated into the choanosome of some spe¬
cies, but are absent from the fibres of all species.
Sponges are intense yellow, green, orange, blue,
and purple in colour.

Ianthella basta (Pallas)

(Fig. la-e, PI. la-b, 2a-b, 3a)

Restricted synonymy.

Spongia basta Pallas, 1766: 309.

Spongia striata Lamarck, 1814: 364.

Ianthella flabelliformis ; de Laubenfels, 1948:
155.

Ianthella basta Gray, 1869: 51; Bergquist,
1980: 498.

Material examined. UAZA 15.3, 15.11,
15.12, 15.14, 15.17, AMZ5091: Aidler’s Bay,
Port Moresby, Papua New Guinea, 10 m, 12
January 1986; NSRC (UPNG) 20: Motupore
Island, Bootless Inlet, Papua New Guinea, 12 m,

22 June 1984; NSRC (UPNG) 24: Buna Motu
Islet, Bootless Inlet, Papua New Guinea, 18 m,

23 March 1985; UAZA 15.30: Apra Harbour,
Guam, 15 m, collector C. Birkland, 26 Novem¬
ber 1985; AM Z5092, UAZA 20.1: Marie-IIelene
Reef, Walinde, West New Britain, Papua New
Guinea, 26 m, 22 October 1991; UAZA 20.6:
Cape Lambert, West New Britain, Papua New
Guinea, 20 m, 18 October 1991;Q66C-4357-A:
Nagada Harbour, Madang, Papua New Guinea,

152

Systematics and biogeography of the genus lanthella in the south-west Pacific

Fig. 1. lanthella hasta (Pallas). A, preserved specimen showing radiating rows of fine, regularly spaced conules,
scale - 2 cm; B, the skeleton is a complex rectangular reticulation of fibre fascicles joined by regularly spaced
secondary connecting fibres, oriented within a single plane. Note that the fascicles consist of up to four fibres aligned
one above the other, perpendicular to the plane of the sponge, scale — 0.5 cm; C, soft tissues are cavernous with large
sac-like, spherical or slightly oval choanocyte chambers set in a sparse mesohyl. Fibres consist predominantly of
bark, scale - 200 pm; D, scanning electron micrograph of the skeleton. Note the short, sharply pointed fibres that
extend outwards from the edges of the fibre fascicles, scale - 300 pm; E, differential growth rates are evident in the
lamella in bands of very closely spaced connecting fibres laid down in new directions, scale - 0.5 cm.

153

P.R. Bergquist and M. Kelly-Borges

20 m, 4 September 1990; UAZA 4.1,6.2: Davies
Reef Lagoon, Great Barrier Reef, Australia, 20
m, 18 August 1982; UAZA 6.1: Darwin Har¬
bour, Northern Territory, Australia, May 1984.

Additional material. MNHN DT578 (82):
Spongia basta Pallas; MNHN DT523 (48):
Spongia striata Lamarck.

Description. Elongate fan or vase, occasion¬
ally with basal fanlets, up to 1500 cm in height
and width, 1-3 mm thick. Sponges are attached
to the substrate at a constricted base which is
thickened and strengthened by crowded fibre
fascicles (Fig. la). Small specimens are semi¬
circular and attached along the base of the
lamella; vase-shaped specimens can be attached
by a solid or circular base. Large specimens are
undulate with waves along the vertical, and
occasionally, the horizontal axis of the sponge
(PI. 1 a,b, 2a,b). The surface is extremely regular
with radiating rows of sharply pointed conules,
0.5-1.0mmhigh, 1.0-2.5 mm apart. Oscules are
0.5-1.0 mm diameter, and inhalant pores are
separated on opposite sides of the sponge la¬
mella; scattered within the depressions between
the fibre meshes.

The sponge is flexible, and the surface is
fleshy but slightly harsh to the touch due to
projecting fibre tips. In life, the external and
internal colouration of the sponge is most com¬
monly bluish violet (P6/6-5/6) in the basal three
quarters of the sponge, gradually becoming
lighter, to greyish magenta (RP7/4) in the outer
regions of the sponge. Sponges can also be mi-
mosayellow (Y7/10), copper green (G7''8-GY7/
8), mandarin orange (YR6/10), opal green (BG5/
6-6/6), or vivid blue (PB7/6). Deep reddish-
purple fibres can be seen through the tissue at the
sponge surface. On exposure to air and in ethanol,
the sponge rapidly turns dark reddish purple
(RP5/2). lanthella basta is oviparous. UAZA
15.12, collected on the 26 January 1986, con¬
tains unfertilised eggs 62-94 pm diameter with
nuclei and prominent nucleoli. No choanocyte
chambers remain intact in this specimen.

Skeleton. The skeleton is a complex rectan¬
gular reticulation of fibres oriented within one
plane, radiating from the base to the margin of
the sponge (Fig. lb, PI. 2a,b). Through the
vertical plane of the lamella, 1-2 mm wide
fascicles composed of two to four fibres 20-100
pm wide, aligned one above the other perpen¬
dicular to the plane of the sponge, run 1-3 mm
apart and parallel to each other (Fig. Id). The
fasciculi are connected at regular intervals of 1

mm by single, or occasionally tracts of two or
three fibres (Fig. Id). Differential growth rates
are evident across the lamella, as bands of very
closely spaced connecting fibres that are laid
down in new directions (Fig. le). Short, sharply
pointed fibres extend outwards from the edges of
the fascicles (Fig. Id). The bark component of
the fibres is strongly laminated and incorporates
spongocytes in concentric arrays, pith makes up
approximately one third of the fibre diameter.

Soft tissue. The soft tissues of lanthella basta
are uniformly cavernous, with slightly ovate
choanocyte chambers 80-100 pm in longest
dimension, and large ramifying exhalant canals
(Fig. lc). There is little mesohyl surrounding the
chambers. A narrow ectosome 21 -73 pm deep is
distinguished from the underlying choanosome
by a lack of chambers, greater tissue density and
collagen reinforcement.

Ecology, lanthella basta is found singly or in
groups, on inshore silty patch and fringing reef
slopes thatexperience good tidal or current flow.
Marie-Helene Reef in West New Britain, and
Aidler’s Bay in Port Moresby, Papua New
Guinea, support a spectacular growth of 2-3
sponges per m 2 , many of which are up to 1.5 m
in height.

All specimens examined were infested with
an unidentified polychaete which occupies the
exhalant canals. A small brown and white striped
apodid holothurian, Synaptula lamperti , has
been recorded in densities of >200 per m 2 on
lanthella basta on the Great Barrier Reef, Aus¬
tralia. These have also been recorded in West
New Britain. Hammond and Wilkinson (1985)
have shown that Synaptula lamperti ingests
organic exudates from the sponge surface and
metabolises these compounds for incorporation
into the body wall.

Remarks. lanthella basta is the best known
and most easily recognised species of lanthella.
This is in part due to its wide distribution in the
Indo-West Pacific, and thus its presence in older
collections, but it is mainly due to the morphol¬
ogy of the sponge. The thin, two dimensional,
fan or vase-like form of lanthella basta is very
distinctive, and the fine reticulation of fibres in
very regular rectangular meshes, is striking.

lanthella basta has been well illustrated and
described by Lendenfeld (1889) and Topsent
(1931). In all specimens examined, there is no
evidence of the development of fibrous exten¬
sions of the basic two-dimensional skeleton.
lanthella basta exhibits the full range of verongid

154

Systematies and biogeography of the genus Ianthella in the south-west Pacific

colouration, but no differences in fibre construc¬
tion or arrangement can be detected between
specimens with differing coloration.

De Laubenfels (1948) synonymised Spongia
striata Lamarck with Ianthella flabelliforrnis.
Spongia striata has been examined and is defi¬
nitely a specimen of /. basta Pallas, having the
fine regular rectangular network characteristic
of the species.

Distribution. Northern Australia, Torres
Straits, Papua New Guinea, Guam, Mascarene
Islands, Indian Ocean.

Ianthella flabelliforrnis (Pallas)

(Fig. 2a-e, PI. 4a-b, 5a-b)

Restricted synonymy.

Spongia flabelliforrnis Pallas, 1766: 380;
Lamarck 1814: 550.

Verongia flabelliforrnis Ehlers, 1870: 11.

Ianthella flabelliforrnis Gray, 1869: 50;
Lendenfeld, 1888: 23; Lendenfeld, 1889: 683
(in part, see I. quadrangulata sp. nov.);
Polejaeff,1884:37; Wilson, 1925:474; Bergquist,
1980: 497.

Material examined. UAZA 15.10, 15.13,
15.16, AM Z5093: Aidler’s Bay, Port Moresby,
Papua New Guinea, 10 m, 12 January 1986;
Q66C-4771 - A: north side of Cumberland Strait,
Wessel Islands, Northern Territory, Australia,
16 m, 14 November 1990; UAZA 29: Aidler’s
Bay, Port Moresby, Papua New Guinea, 15 m, 8
January 1984;Q66C-4532-R: Mananouha(Lion)
Island, Bootless Inlet, Papua New Guinea, 22 m,
15 September 1992; NTM Z1410: north-east of
Port Hedland, Station NWS 17, 80 m, 19 April
1983; NTM Z908: Dudley Point, Darwin, North¬
ern Territory, Australia, Station EP8, 10 m, 31
August, 1982, collector J. N. A. Hooper; NSRC
(UPNG) 114. Q66C-4532-R: Mananouha(Lion)
Island, Bootless Inlet, Papua New Guinea, 22 m,

7 March 1985; NTM Z689: West Buccaneer
Archipelago, Western Australia, Station DON
19, 35 m, 28 April 1982; NTM Z850: Channel
Island, Darwin, Northern Territory, Australia,
Station CI3,12-13 m, 20 August 1982, collector
J. N. A. Hooper; NTM Z1072: Fish Reef, Bynoe
Harbour, Northern Territory, Australia, Station
FRI, 10-12 m, 24 November 1982, col lector J. N.
A. Hooper; BMNH 1930.8.13.164: Great Bar¬
rier Reef, Australia; BMNH 1885.8.8.1: south of
New Guinea,9° 59’S, 139°42’E. Station 188,56
m, 10 September 1874.

Additional material. AM - G9980Lord Howe
Island, Australia; AM - G9975: Lord Howe
Island, Australia; AM - G9981: Manly Beach,
Australia; AM - G8905: Port Jackson, New'
South Wales; NTM Z2557; MNHN DT534,
DT3448, DT3449: Spongia flabelliforrnis
Lamarck.

Description. Symmetrical single or
bilamellate fan (Fig. 2a, PI. 4a,b), up to 1000 cm
high, 10-15 mm thick in the centre, 1-3 mm on
the outer edge of the lamella. Small sponges are
elevated by a fibrous stalk. In large sponges the
base is broad and thickened by robust fibre
fascicles. The surface is raised into narrow ir¬
regular ridges 2-5 mm high, except in the outer
40-50 mm of the lamella. Oscules, 1-2 mm wide,
are located on one face of the sponge. The
texture isharsh. In life the external and internal
colourof the sponge is most commonly greenish
yellow (GY8/10), occasionally vivid blue (PB7/
6) with a whitish cast due to the presence of sand
grains in the dermal tissue. Reddish purple
fibres (RP3/10) are clearly seen in the surface of
the sponge. On exposure to air or ethanol, the
sponge is dark reddish purple. Ianthella
flabelliforrnis is oviparous, eggs are present in
specimen Q66C 4771-A, collected on 10 No¬
vember 1990. Eggs are 15-73 pm in diameter,
ovoid, and grouped in clumps within the
choanosome (Fig. 2b, PI. 5a,b).

Skeleton. The skeleton is a semi-regular re¬
ticulation of fibres, 120-500 pm thick, oriented
in one plane and radiating from the base to the
margin of the sponge (Fig. 2d,e). Primary fibres
are loosely fasiculated and interconnected by
smaller fibres to form irregular polygonal meshes
2-5 mm wide and 5-7 mm high; variants of this
basic pattern occur. In specimen NTM Z1410,
the reticulation is irregular and anastomosing,
with primary and secondary fibres barely distin¬
guishable. In NSRC (UPNG) 114, the secondary
fibres dominate the skeleton and coalesce in
places to form a fibrous plate.

Outgrowths from the plane of the lamella are
regularly spaced 5-10 mm apart and are com¬
posed of fasciculate extensions of the primary
fibres that coalesce at the apex to form a round
flattened knob up to 5 mm in diameter; the apex
of the knob is minutely conulose (Fig. 2c,e).
Several adjacent bosses may coalesce to form a
surface pattern of meandering, often intercon¬
nected ridges. Knobs are more pronounced to¬
wards the centre and lower portions of the
sponge, and on the poral face.

155

P.R. Bergquist and M. Kelly-Borges

Fig. 2. lanthella flabeUiformis (Pallas). A, preserved specimen showing narrow irregular interconnected ridges, scale - 3 cm;

B, the choanosome is cavernous, the choanocytes are grouped within a thick collagenous mesohyl. Fibres are thick, the pith
comprising approximately 10 % of the volume of the fibre. Note oocytes grouped in choanosome, scale - 300 pm;

C, scanning electron micrograph of fibrous skeletal outgrowths. These are composed of fasciculated extensions of the primary
fibres that coalesce at the apex to form a round flattened knob, the apex of which is minutely conulose, scale - 100 pm; D,
scanning electron micrograph of the skeleton which is a semi-regular reticulation of fibres oriented in a single plane, radiating
from the base to the margin of the sponge. The primary fibres are loosely fasciculated and interconnected by smaller fibres to
form irregular polygonal meshes, scale - 200 pm; E, adjacent fibrous outgrowths may coalesce to form a surface pattern of
meandering ridges, scale - 0.5 cm.

156

Systematics and biogeography of the genus lamhella in the south-west Pacific

The bark component of the fibres is strongly
laminated and incorporates spongocytes in con¬
centric arrays, pith comprises approximately 10
% of the volume of the fibre.

Soft tissue. The choanosome is cavernous,
but choanocyte chambers lie in a moderately
thick mesohyl (Fig. 2b). Theectosomal region is
heavily reinforced with collagen and is 100-170
pm deep. Sand grains are scattered throughout
the ectosome and outer choanosome.

Ecology. The sponge occurs on coral reef
slopes in oceanic or inshore waters which expe¬
rience good current movement. An unidentified
polychaete is almost always present in the exha-
lant canals.

Remarks. Examination of fresh specimens of
Ianthella flabelliformis from Papua New Guinea
and Australia confirm the major features of fibre
construction and the choanosome as described
by Lendenfeld (1889) and Pol6jaeff (1884). This
species differs from I. basta in having a much
less regular skeletal arrangement, as well as in
having complex fasciculated fibre extensions
perpendicular to the two dimensional plane of
the sponge. In /. basta , these extensions are
short sharp spikes, and the sponge lamella is
much thinner as a result. The surface of /.
flabelliformis, is much less regular than that of
I. basta and the choanosome can contain sand
and detritus.

Our observations of the architecture of surface
extensions of Ianthella flabelliformis differ in
detail from the descriptions given by Lendenfeld
(1888), Potejaeff (1884) and Wilson (1925). The
fibres perpendicular to the surface of the sponge
have been described as being “ramified in a
dendritic manner’’ emanating from the junction
of the primary radial and secondary connecting
fibres (Lendenfeld 1888). These fibres in fact
arise at irregular intervals along the primary
radial fibres and form blunt fasciculations.

There are differences in colouration and the
degree of flexibility of the sponge, between
specimens of Ianthella flabelliformis described
here and those described in older literature.
The rigidity of the skeleton is dictated by the
robustness of the individual fibres, the degree
of fasciculation of the primary and secondary
fibres, and the presence of surface extensions.
The amount of sand incorporated in the sponge
tissue will also affect the pliability of the sponge.
Bergquist (1980) noted considerable variation
among specimens labelled as this species in the
London Natural History Museum collections.

In particular, some specimens had very thick
basal fibres. Many of these specimens, mainly
dry, can be assigned to two of the new species
described below. Lendenfeld’s specimen labelled
as Ianthella flabelliformis (BMNH 1886.7.8.13)
from Port Jackson has very large fibres and is
the holotype of /. quadrangulata, described be¬
low.

Ianthella flabelliformis is common in south¬
ern Papua New Guinea and along the eastern
and northern coasts of Australia, and is reported
to extend as far south as Port Jackson. This
southern record rests on an identification by
Lendenfeld (1889) of a specimen collected by
Ramsay. It was only after publication of this
work that Lendenfeld realised that another spe¬
cies, I. quadrangulata, could be recognised. It is
probable therefore that the true southern limit of
I. flabelliformis is the southern Great Barrier
Reef. This species is most easily recognisable in
the field by its frequently bilamellate construc¬
tion, harsh texture, and irregular embossed sur¬
face.

Distribution. Western, northern, eastern and
south-eastern coasts of Australia, Torres Straits,
southern Papua New Guinea.

Ianthella labyrinthus sp. nov.

(Fig. 3a-c)

Type material. HOLOTYPE - NTM Z691:
west of Buccaneer Archipelago, Western Aus¬
tralia, Station DON 19, 35 m, 28 April 1982.

Description. Highly symmetrical circular fan,

140 mm wide and 165 mm high, attached to the
substrate by a thick flattened stem 20 mm in
diameter and 50 mm high (Fig. 3a). The lamella
is 12-14 mm thick basally and 2-3 mm thick at
the margin. The surface is thrown into meandrine
ridges, which are rounded apically and minutely
conulose. Ridges are 1-4 mm wide and 2-5 mm
high on the face, and have a roughly concentric
arrangement on the oscular face. The outer 7-15
mm of the fan is regularly and minutely conulose.
The lamella is flexible, Finn and just compress¬
ible, the texture rough. In life, the internal and
external colouration is greenish yellow (GY8/
10), with reddish purple fibres visible through
the surface tissue. In ethanol the tissue is deep
reddish purple (P3/8).

Skeleton. Primary fibres are irregularly
fascicu lated in groups of three to five fibres, each
70-200pm in diameter, interconnected by slightly
finer secondary fibres to form small irregular

157

P.R. Bergquist and M. Kelly-Borges

Fig. 3. lanthella labyrinthus sp. nov. A, preserved specimen (holotype, NTM Z69 1 ) showing the meandrinc, interdigitating,
rounded minutely conulose ridges, scale - 2 cm; B, choanocyte chambers are grouped in a thick mcsohyl, the cortex is heavily
reinforced with collagen, and sand and spicular debris is scattered within the cortex. Fibres are fine and consist predominantly
of bark, scale - 200 pm; C, the skeleton consists of fine irregularly fasciculated fibres that are connected by finer secondary fibres
to form small polygonal meshes. Complex irregular outgrowths consisting of fine anastomosing fibres extend outward from the
sponge lamella, coalescing to form a labyrinthine surface pattern, scale = 300 pm.

158

Systematics and biogeography of the genus Ianlhella in the south-west Pacific

polygonal meshes 1 -2 mm wide and 5-7 mm high
(Fig. 3c). At intervals of 5-10 mm along the
fascicular columns, the fascicle broadens and
complex outgrowths arise and these coalesce to
form elaborate surface ridges. The ridges are
arranged approximately concentrically toward
the sponge margin, or in a labyrinthine pattern
towards the centre of the sponge. The surface
extensions are made up of fine irregularly
anastomosing fibres20-50 pm diameter(Fig. 3c).
The outer 10-15 mm of the sponge margin is very
thin and the skeleton in that region is more regu¬
lar than at the centre of the sponge lamella.

Individual fibres are up to 200 pm thick and
have a very reduced pith. It makes up 5-10 %
only of the fibre diameter. The bark is loosely
laminated with spongocytes incorporated in con¬
centric arrangement.

Soft tissues. Choanocyte chambers are
grouped in -a dense mesohyl (Fig. 3b). The
ectosome is heavily reinforced with collagen
and is 70-120 pm deep. Sand and spicular debris
is scattered within the ectosome.

Ecology. The sponge was found attached to a
sand-covered rock surface at 35 m.

Etymology. The surface patterning of the
sponge is like a maze.

Remarks, lanthella labyrinthus is compara¬
ble to I. flabelliformis in that the basic two-
dimensional skeleton of the sponge is supple¬
mented with fibrous extensions that form an
elaborate surface pattern of ridges and knobs.
The two species are also greenish yellow and
both have a stalked, roughly circular lamella.
However, there are differences in skeletal ar¬
rangement and resultant surface patterns be¬
tween the two sponges. The surface of /.
flabelliformis has very firm squat knobs that
render the surface harsh and irregular, while the
surface extensions of /. labyrinthus form soft
microconulose concentric ridges. The fibres oil.
labyrinthus are much finer than those of /.
flabelliformis and the pith component of the
formerspecies is greatly reduced in comparison.
lanthella labyrinthus has a much thicker body
than I. flabelliformis and this permits develop¬
ment of a dense mesohyl.

The characteristic features of lanthella
labyrinthus are the surface patterning and the
fine, slightly irregular rectangular reticulation
at the outer margin. The skeleton and resultant
surface pattern of the outer margin resembles
the surface of/, basta, although the mesh is not
as regular and the radial fascicles are wider.

lanthella quadrangulata sp. nov.

(Fig. 4a-c, PI. 6a-b)

Type material. HOLOTYPE - BMNH
1886.7.8.13: Port Jackson, New South Wales
(misidentified as lanthella flabelliformis by R.
von Lendenfeld).

Additional material. UAZA 25: Fairlight
Harbour, Sydney, New South Wales, Australia,
2 February 1964; AM G8906: Port Jackson, New
South Wales; FN1334: Port Jackson, New South
Wales; BMNH 1844.9.13.32: Illawarra, New
South Wales, Australia, presented by J. B. Jukes
Esq, collected by H. M. S. Fly, BMNH 39.2.4.1:
Lord Howe Island, Australia, presented by L. B.
Moore; BMNH 1925.11.1.883: Manly Beach,
Australia; BMNH 1938.5.2.10: Western Aus¬
tralia, presented by D. L. Serventy, Perth; NTM
Z 1784B: west side of Russell Island, Frankland
Island Group, Western Australia, 23-26 m; QM
G303784; Long Reef, Sydney, New South Wales,
Australia, 50 m, collector D. Roberts, 16 August
1993;QM G301211: Pig Is..Port Stephens, New
South Wales, Australia, 10 m, collector M.
Garson, 1989. QM G304088: Lazarette Gutter,
Peel Island, Moreton Bay, Queensland, Aus¬
tralia, 4 m. Collector N. Coleman, 6 October
1993; QM G300027: Outer Gneerings Shoals,
off Mooloolaba, Queensland, Australia, 20 m,
collector J. N. A. Hooper, 10 December 1991;
QMG303942: JewShoals, Noosa Head, Queens¬
land, Australia, 20 m, collector J. N. A. Hooper,
9 February 1994.

Description. The sponge is an elongate fan,
10-16 cm high and wide, and 3-5 mm thick with
basal fanlets. It has a slightly constricted base of
attachment which is strengthened by thick, in¬
terlocking primary fibres. The surface is smooth
and the lamella is flexible across the horizontal
axis only. In life the external and internal
colouration is mimosa yellow (Y8/12). On expo¬
sure to air, and in ethanol, the sponge turns dark
reddish purple (RP5/2).

Skeleton. The skeleton is a simple reticula¬
tion of robust vertical fibres 0.5-2.5 mm in
diameter, anastomosing to produce a net-like
skeleton. The shape of the meshes is elongate
and polygonal, measuring up to 20 mm long by
7 mm wide. Finer secondary fibres, 0.2-0.5 pm
wide are common, irregularly distributed, and
are oriented diagonally across the fan. Fibre pith
occupies more than 50 % of the fibre diameter
and the bark is laminated, charged with
spongocytes in concentric array.

159

P.R. Bergquist and M. Kelly-Borges

Plate 1. A, Ianthella basta, Papua New Guinea, in situ, specimen 1 .0 m high; B, lanthella basta, Papua New Guinea, in situ,
specimen 1.5 m high.

160

Systematics and biogeography of the genus lanthella in the south-west Pacific

Plate 2. A, lanthella basta, Papua New Guinea, close up of regular rectangular skeleton in the living sponge, natural size;
B, lanthella basta, Papua New Guinea, close up of undulating body structure, natural size.

161

P.R. Bergquist and M. Kelly-Borges

Figure 4. Ianlhella quadrangulata sp. nov. A, preserved specimen (holotype, BMNH 1886.7.8.13)
showing widely spaced sharply pointed surfaceconules, scale - 2 cm; B, preserved specimen showing
the large anastomosing primary Fibres interconnected by smaller secondary fibres, scale - 1 cm; C, the
skeleton is a simple reticulation of robust vertical fibres anastomosing to produce a net-like skeleton
with elongate to quadrangulate meshes, scale = 1 cm.

162

Systematics and biogeography of the genus lanthella in the south-west Pacific

Soft tissue. Choanocyte chambers are grouped
in a strongly collagenous mesohyl. A large por¬
tion of the tissue volume is taken up by broad
tracts of collagen that run vertically through the
sponge, and surround the fibres.

Ecology. lanthella quadrangulata is found
predominantly within harbours and in turbid
coastal waters, down to 50 m.

Etymology. The species name describes the
shape of the meshes produced by anastomosis of
the huge vertical fibres in this species.

Remarks. The major characters that distin¬
guish lanthella quadrangulata from other spe¬
cies of lanthella are the detail of skeletal struc¬
ture and the extremely collagenous mesohyl.
The skeleton is a network of extremely thick
anastomosing fibres which lack complex lateral
outgrowths. The fibres in 1. quadrangulata are
very thick, and appear to be hollow on drying
due to the substantial pith volume.

lanthella quadrangulata is the southernmost
species of this genus and is known to extend only
as far north on the east coast of Australia as
Hervey Bay in Queensland where it overlaps
with t.flabelliformis at its southern limit around
the Fraser Island and Tweed Heads regions (J.
N. A. Hooper, pers. comm. 1995). Both species
have a similar depth distribution.

Bergquist (1980) noted that in the collections
labelled as lanthella flabellifonnis in London’s
Natural History Museum and the Australian
Museum, there were dry specimens that had
unusually large fibres. In particular, a group of
specimens labelled lanthellaflabelliformis (AM
G9976) from New South Wales contained a
number of fragmentary specimens of lanthella
from the Thetis collection from Lord Howe
Island, all of which had large fibres. Although
positive identification was not possible due to
the state of the specimens, it is likely that they
represent additional records of lanthella
quadrangulata.

In an unpublished manuscript held by the
Natural History Museum, London, Lendenfeld
described specimens of lanthella from various
southern New South Wales collections that are
very similar to the specimens described above.
Lendenfeld suggested that the New South Wales
specimens be given the species name
quadrangulata. which accurately reflects the
shape of the skeletal meshes in his specimens
and those specimens examined in this study. We
have chosen to formally adopt the species name
quadrangulata, and have designated a further
good Lendenfeld specimen labelled /.

flabelliformis (BMNH 1886.7.8.13) as the
holotype of /. quadrangulata.

lanthella reticulata sp. nov.

(Fig. 5a-e, PI. 3b)

Dendrilla aerophoba', Pulitzer-Finali 1982:136.

Type material. HOLOTYPE - AM Z5094:
Aidler’s Bay, Port Moresby, Papua New Guinea,
10 m, 12 January 1986 (UAZA 15.15).

Additional material. UAZA 15.4, 15.8, AM
Z5095: Aidler’s Bay, Port Moresby, Papua New
Guinea, 10 m, 12 January 1986; UAZA 30:
Aidler’s Bay, Port Moresby, Papua New Guinea,
15 m, 8 January 1984; NSRC (UPNG)26: Buna
Motu Islet, Bootless Inlet, Papua New Guinea,
18 m, 22 August 1984; UAZA 15.GBR: Davies
Reef, Great Barrier Reef, Australia, 10 m, 30
October 85; NSRC (UPNG) 70: Mananouha
(Lion) Island, Bootlesslnlet, Papua New Guinea,
20 m, 17 August 1985; AIMS PA 23: Pandora
Reef, Great Barrier Reef, 12 m, 28 May 1984,
collector C. R. Wilkinson; UAZA 31,32,33,34:
Davies Reef Lagoon, Great Barrier Reef, 20 m,
18 August 1982, collectorC. R. Wilkinson; AM
Z4113 (Roche FN1915): Russell Is., 12-20 m;
AM Z4137: Escape Reef, Great Barrier Reef, 20
m; BMNH 1993.11.19.1 (0CDN 2048-L): Kar

Karlsland, north of Madang, Papua New Guinea,

20 m, collector P. L. Colin, CRRF, 19 November
1993.

Description. Semi-circular or irregularly
shaped fan, 10-25 cm high and wide, and 5-10
mm thick, frequently prostrate or curled over,
attached along the entire length of the lamella
(Fig. 5a). The surface is raised into prominent
well spaced multiple conules, 5-7 mm high, and
3-8 mm apart, the surface is spikey but very
smooth and fleshy between the conules. The
texture is elastic and compressible. Oscules 0.5-
1.0 mm wide, are regularly distributed on one
surface of the sponge. In life the external and
internal colouration is mandarin orange (YR6/

10), mimosa yellow (Y8/12) or bluish violet (P6/
6-5/6). On exposure to air, and in ethanol, the
sponge is dark reddish purple (RP5/2).

Skeleton. The skeleton is a simple reticula¬
tion of uniform anastomosing fibres, 0.5-1.5 pm
in diameter, in a net-like arrangement (Fig. lc).
The primary fibres are zigzagged in the vertical
plane, and, through the plane of the lamella, the
meshes formed are diamond-shaped and 10-15
mm long by 1-5 mm wide. Fine short secondary
fibres 0.2-0.5 pm wide, are oriented diagonally
across the fan but only connect between a few

163

P.R. Bergquist and M. Kelly-Borges

Plate 3. A, lanthella basta, Guam, in situ, specimen 1 m high; B, lanlhella reticulata, Papua New Guinea, close up view of
fleshy surface and irregular skeletal pattern, natural size.

164

Systematics and biogeography of the genus hmthella in the south-west Pacific

Plate 4. A, lanthellaflabelliformis, Papua New Guinea, in situ, specimen 40 cm high; B, lanlhcllajlabelliformis, Papua New
Guinea, in situ, specimen 30 cm high.

165

P.R. Bergquist and M. Kelly-Borges

Fig. 5. lanthella reticulata sp. nov. A, preserved specimen showing the prominent, well-spaced multiple conules, scale
= 1.5 cm; B, dendritic spikes extend outwards from zigzagged primary fibres, evident in this lateral view of the skeleton.
The oscular face of the sponge is on the right, scale - 0.5 cm; C, scanning electron micrograph of the dendritic skeletal
outgrowths, scale = 100 pm; D, the skeleton is a simple reticulation of uniform anastomosing fibres that is net-like in
arrangement. Note the fine diagonal fibres, scale - 0.5 cm; E, choanocyte chambersare large, spherical or elongate and
arc grouped in a mesohy I heavily reinforced with collagen. Pith occupies approximately 50 %of the fibre width, and fibres
are sheathed with collagen, scale = 200 pm.

166

Systematics and biogeography of the genus lanthella in the south-west Pacific

adjacent fibres. These fibres are patchy in occur¬
rence. Dendritic spikes extend outwards from
the zigzagged primary fibres; this is evident in
the lateral view of the skeleton (Fig. 5c,d), and
are more pronounced on the poral face of the
sponge. The fibre pith occupies up to 75 % of the
fibre width and the bark laminae are charged
with spongocytes in concentric array. Often
multiple pith elements are surrounded by a
common bark layer.

The rigidity of the skeleton varies with the
robustness of the fibres, and the degree of their
interconnection. Sponges also vary in the degree
of lateral ornamentation of the fibre.

Soft tissue. Choanocyte chambers are large,
oval or elongate, and the mesohyl is heavily
reinforced with collagen. Collagen surrounds
the fibres as a sheath. The ectosome on the poral
face of the sponge is 147-367 pm deep, strongly
collagenous, and honeycombed with inhalant
apertures 62-135 pm wide. The ectosome on the
oscular surface is compact, 50-170 pm deep and
also heavily reinforced with collagen.

Ecology. lanthella reticulata is moderately
common, and found on inshore fringing reef
slopes. Specimens may be heavily infested with
a commensal barnacle, Acasta porata, which
becomes embedded in the sponge body and
attached basally to the fibrous skeleton of the
sponge.

Etymology. The skeleton is reticulated like
the mesh of a net.

Remarks. lanthella reticulata is closely re¬
lated to I. quadrangulata, which also has
anastomosing fibres, an unstalked body and a
choanosome that is heavily reinforced with col¬
lagen; both possess fibres that are similarin size,
structure, and arrangement. They differ in the
shape of the skeletal meshes, however, which
are zigzagged and polygonal in/, reticulata, and
flattened and polygonal in I. quadrangulata.
lanthella reticulata possesses well developed
dendritic spikes in a plane perpendicular to the
surface of the sponge; these are absent in /.
quadrangulata. Secondary fibres are quite vari¬
able in their distribution within /. reticulata,
while in I. quadrangulata the secondary fibre
network is better developed. The two species are
geographically separated: /. quadrangulata oc¬
curs in southern Australian waters, and /.
reticulata is found in north-eastern Australian
waters, southern Papua New Guinea, and occa¬
sionally on the north coastof Papua New Guinea.

Pulitzer-Finali (1982) identified a fan-shaped
yellow sponge from the Great Barrier Reef as

Dendrilla aerophoba Lendenfeld. Dendrilla
aerophoba was originally described from Port
Philip in Southern Australia, and is described as
having strange grooved fibres, and a highly
structured cortical armour of clionid chips
(Lendenfeld 1889). These novel features are not
mentioned in Pulitzer-Finali’s brief recent de¬
scription of the sponge. The Great Barrier Reef
specimen is described as being a 10 mm thick
fan with a dendritic, anastomosing skeleton of
dark laminated fibres, with fibrous offshoots
perpendicular to the main skeleton. There is no
doubt from this information and from photo¬
graphs of the specimen that it is lanthella
reticulata. Dendrilla aerophoba Lendenfeld is
unrecognisable on the basis of present collec¬
tions except as belonging to the Ianthellidae.

Hyatt (1875) described lanthella concentrica
from the Fiji Islands, a sponge that was “prob¬
ably irregularly frondose” and 3-6 mm thick.
The skeleton consisted of what appear to be thick
fibres 1-3 mm in diameter, “compound and
ornamented with thick short spines and minute
wart-like prominences”. Hyatt also noted that
“the fibres are almost always double, contained
within common layers of homy material and
externally appearing as one stem” which is
reminiscent of the multiple piths seen in lanthella
reticulata fibres. The skeletal mesh is described
as being “quite irregular in size and shape,
varying from 12 mm to only 5 mm in length”. It
is possible that /. concentrica is synonymous
with lanthella reticulata, but there are no speci¬
mens, other than the type which has been de¬
clared unrecognisable, available to permit pre¬
cise identification.

Among species ol lanthella, the skeletal struc¬
ture of lanthella reticulata and I. quadrangulata
come closest to that of Anomoianthella, but the
latter sponge has a definite and well developed
three dimensional interlocking skeletal struc¬
ture. Lendenfeld (1888, 1889) placed Western
Australian specimens of a thick-walled, cup¬
shaped sponge with clumped oscules and huge
fibres in lanthella concentrica (BMNH
1886.2.19.13). These specimens belong to
Anomoianthella (Bergquist 1980).

DISCUSSION

The range of morphologies known within the
genus lanthella has been extended with the
description of three new species and by
redescription of lanthella flabelliformis and /.

167

P.R. Bergquist and M. Kelly-Borges

Plate 5. A, lanthellaflabelliformis. Photomicrograph to show fibre structure, eurypylous choanocyte chamber structure and
developing eggs x 200; B, lanthella flabelliformis. Photomicrograph to show detail of egg structure x 600.

168

Systematics and biogeography of the genus Ianthella in the south-west Pacific

Plate 6. A, Ianthella quadrangulata. Holotype. Low power photomicrograph to show strong ectosomal development and fibre
structure x 150. This specimen has been preserved since 1889 and the choanosomal tissue has deteriorated;
B, Ianthella quadrangulata. Mooloolaba, Queensland, Australia, in situ, specimen 70 cm high.

169

P.R. Bergquist and M. Kelly-Borges

Table 1. Characters and character states of lanthella. ' Character states are unique to the species listed. 2 Ianthellid characters
13 and 14, and 15 do not apply to the outgroups, as Psammaplysilla purpurea and Pseudoceratina durissima do not possess
reticulate skeletons, and their skeletons are not compressed into two dimensions as they are in lanthella. Outgroup character
states 13, 14 and 15 were thus coded as absent. Character states in italics are unique to the outgroups. J Although species of
lanthella are relatively uniform in thickness, the surface is frequently ornamented with ridges, knobs and conules, and specimens
areofien thicker in thecentreofthe lamella. Character statesin character 1 are thus defined as the maximum thickness, including
ornamentation, in thecentre of the lamella. 4 It isdifficult to distinguish between the various hues of yellow that are found within
lanthella andtheaplysinellid outgroups. Yellow pigmentation ranges from golden or mimosa yellow to lemon in lanthella and
Pseudoceratina , to greenish yellow or yellow tinged with bright green in Psammaplysilla and lanthella. Only clearly defined
colours are included in the character list. A. popae - Anomoianthella ; I. - lanthella.

MORPHOLOGY

1. Thickness of lamella 3

a. 1-3 mm: /. hast a'

b. 5-10mm

c. 10-15mm

d. >20mm: A. popae'

e. non-lamellate

2. Attachment to substrate

a. distinct stalk or basal constriction of fasciculated
primary fibres

b. stalk absent with only a slight restriction of
sponge base

3. Morphology

a. single fan

b. bilamellale: l.flabelliformis 1

c. principal fan plus smaller additional fans

d. knob-shaped

e. complex encrusting

COLOURATION 4

4. blue violet

a. present

b. absent

5. mandarin orange

a. present

b. absent

6. vivid blue

a. present

b. absent

SKELETAL STRUCTURE

7. Body compressed in a two planes

a. present

b. absent

8. General organisation

a. fibro-reticulate

b. dendritic

9. Fibretsoft tissue volume

a. fibre dominates

b. sparse fibre

FIBRE CONSTRUCTION

10. Proportion of bark in fibre

a. significant component of fibre

b. absent or reduced to patches

11. Spongocytes in bark

a. present;

b. absent

SKELETAL ARCHITECTURE

12. Primary fibres

a. fasciculated in a plane perpendicular to the
plane of the fan. very regular: /. basta'

b. irregular non-aligned fascicles

c. single and anastomosing

d. single and dendritic

SKELETAL ARCHITECTURE (cont.)

13. Mesh shape 2

a. extremely regular and rectangular: /. basta'

b. vaguely rectangular to elongate

c. anastomosing primaries form quadrangulate to
oval meshes

d. absent

14. Mesh size (av. Hgt x Wdth) 2

a. 1x3mm: /. basta'

b. 7x3 mm

c. 20x10mm

d. >20mmxl0: A. popae'

e. absent

15. Outgrow ths on the two-dimensional skeleton 2

a. boss-like fasciculated outgrowths that render the
sponge surface ridged and knobbed

b. dendritic outgrowths forming multiple surface
conules: I. reticulata'

c. sharply pointed single spikes forming a very
regular conulose surface: I. basta'

d. absent

16. Maximum fibre thickness

a. 150um

b. 500um

c. 2500um

d. >2500um: A. popae'

SOFT TISSUES

17. Density stratification

a. cavernous; chambers set in sparse mesohyal:

/. basta'

b. chambers grouped in a moderately dense matrix;
collagen evenly distributed.

c. choanosome heavily infiltrated with collagen,
often in tracts

d. heavily infiltrated with collagen, extremely
dense

18. Average cortex depth

a. 70um: /. basta'

b. I20-I70um

c. >300um

19. Cortex structure

a. collagen reinforced

b. fibrous cuticle: A. popae'

20. Choanocytes

a. curypylous, sac-shaped

b. eurypylous, elongated and occasionally
branched: A. popae'

c. diploidal, spherical and small

BIOCHEMISTRY

21. Aplystanc sterols

a. present ;

b. absent

170

Systematics and biogeography of the genus lanthella in the south-west Pacific

Table 2. Character state matrix for lanthella. Characters and states are described in Table 1. When a character state is unknown
for a particular species, the character is coded with a question mark. Asterisks indicate outgroups. Data for outgroups
Psammaptysilla purpurea and Pseudoceratina durissima were obtained from Kelly-Borges and Bergquist (1988), Bergquist
(1965), and Bergquist (1980), respectively. Note that character 19 is uninformative in a cladistic sense, but it is included here
for completeness of the character set and for potential information in future work. According to parsimony criteria, a character
is informative only if, for the given set of taxa, at least two taxa differ in their assigned character state from the remainder of the
set. This is a putative synapomorphic character. Uninformative characters such as the autapomorphic character 19 occur when
a character state is different in only one taxon in the entire group for that character. These characters do not illuminate ingroup
relationships but are indicative of evolutionary rate or ''uniqueness” of the taxon (Swofford, 1991b).

Species

Characters 1-21

Order Verongida

Family Ianthellidae

lanthella basta

lanthella labyrinthus

lanthella flabelliformis

lanthella reticulata

lanthella quadranpulata

Anomoianthella popeae

Family Aplysinellidae

Pseudoceratina durissima

Psammaplysilla purpurea

basta. Traditional diagnostic characters for
lanthella have been re-evaluated and details of
skeletal architecture, fibre construction, and pig¬
mentation have been given consideration. In
particular, we emphasise the pattern of reticula¬
tion of the primary radiating fibres, the nature of
the secondary connecting fibres, and the devel¬
opment of outgrowths of the skeleton from the
plane of the lamella, in redefinitions of taxo¬
nomic characters and thus, distinction between
species.

A phylogenetic analysis of morphological
characters (Tables 1 -2) described here, was car¬
ried out to examine inter-species relationships
within lanthella. Additionally, data for
Anomoianthella popae Bergquist was included
within the analysis, in order to explore further
the position of this genus within the Family
Ianthellidae. There is no evidence of compres¬
sion of the skeleton into a single plane in
Anomoianthella, as there is in lanthella, and the
fibrous skeleton forms an irregular anastomosing
reticulum (Bergquist 1980). The choanocyte
chambers of Anomoianthella are eurypylous but
elongate and occasionally branched.

In order to obtain a directed phylogenetic
analysis, two members of the verongid family
Aplysinellidae, Pseudoceratina durissima Carter
and Psammaplysilla purpurea Carter, were cho¬
sen as outgroups. The characters that separate
these aplysinellid species from lanthella are
possession of a sparse, three dimensional
dendritic skeleton composed of fibres that do not
contain cellular elements, and in which the bark

component is significantly reduced or absent.
These aplysinellid genera also contain aplystane
sterols which are absent in the Ianthellidae
(Bergquist et al. 1991). Contention over the
correct family name to apply to our outgroup
genera has been addressed by Bergquist (1995)
and the name Aplysinellidae is upheld.

The analysis was carried out using PAUP
Version 3.0 (Swofford 1991a) (Tables 2-3) as
described i n Bergquist and Kelly-Borges (1991).
The data were unordered, unweighted, and the
branch and bound option wasused toensure that
all minimum length trees were obtained. Two
equally parsimonious trees were obtained for the
data set, each of length 53, and each with a high
consistency index (Cl) of 0.868. These trees
differed only in their placement of lanthella
quadrangulata and lanthella reticulata in rela¬
tion to each other. Hypothetical relationships
within lanthella and within the Family
Ianthellidae are presented in Figure 6.

The phylogenetic tree indicates that the
Ianthellidae is monophyletic with respect to the
Aplysinellidae, as represented by the outgroups.
The major synapomorphies of the Ianthellidae,
or the changes that occur between nodes 1 and 2
in Figure 6, are the possession of a reticulate
rather than a dendritic fibrous skeleton (charac¬
ter 8B->A) in which fibres have a significantly
amplified bark component (10B->A) and wh ich
contain spongocytes in annuli (11B->A). Fibre
dominates in volume over soft tissue in
Ianthellidae (9B->A). The choanocyte cham¬
bers in the Ianthellidae are eurypylous and sac-

171

P.R. Bergquist and M. Kelly-Borges

shaped (20C->B) and these sponges lack
aplystane sterols (21A->B).

Anomoianthella is placed with Ianthella in
the Ianthellidae as it shares features of fibre and
soft tissue construction. It also has a similar
skeletal structure to Ianthella quadrangulata and
I. reticulata, although the construction is three
dimensional. Anomoianthella differs, however,
from the genus Ianthella in that the body is not
compressed into a thin fan (7B->A), and the
sponge body is not stalked or basally constricted
(2B->A). A fibrous cuticle (19A->B) is unique
to Anomoianthella and choanocyte chambers are
elongated and occasionally branched (20A->B).

Ianthella is monophyletic, with /. basta, 1.
flabelliformis and /. labyrinthus being more
closely related to each other than to /. reticulata
and I. quadrangulata, which are situated basally
on the tree above Anomoianthella popae. Major
characters that differentiate species within the
Ianthella group include the arrangement of the
primary fibres, the mesh shape and size, and the
nature of the surface skeletal outgrowths. The
skeletal factors that separate I. quadrangulata

and I. reticulata (nodes 3 to 4) from other
species are essentially the differences in the
nature of the surface ornamentation of the
sponges. /. reticulata has dendritic outgrowths
of the skeleton, while in /. quadrangulata these
are limited to occasional small spikes.
Synapomorphies for /. basta, I. flabelliformis,
and /. labyrinthus (nodes 4 to 5) include a
complex development of the skeleton with pri¬
mary fibre fascicles (12C->B) in 1. flabelliformis,
1. labyrinthus and /. basta, and smaller more
regular meshes (13C->B, 14C->B) in /.
quadrangulata and I. reticulata. In I.
quadrangulata and I. reticulata, the primary
fibres form a loose anastomosing network, con¬
nected by small diagonally oriented fibres. Fi¬
bres in /. basta,1 .flabelliformis and I. labyrinthus
are also much thinner (16C->A). Tissue density
is reduced in these later three species, and there
is less collagen reinforcement of the matrix. The
choanosome of Ianthella ranges in density from
cavernous in l. basta, I. flabelliformis and I.
labyrinthus, to heavily reinforced with collagen
in I. quadrangulata and /. reticulata.

Table 3. Data from phylogenetic analysis of species of
Ianthella. The characters given in Table 1 are listed with the
number of character states. The column headed Status refers
to the a priori definition of the character as questionable
(characters that arc susceptible to coding errors by misinter¬
pretation), or well defined (characters that can be assigned
with confidence to discrete character states, see Bergquist and
Kelly-Borges, 1991). The character states of the basal branch
of all Ianthella species is given along with the pleisiomorphic
states of the ingroup Ianthella.

Character

states

Number

Status

Consistency

value

Basal branch
of states

for Ianthella

1.000

0.750

A/C

1.000

A/B

0.500

1.000

0.750

1.000

Ianthella basta
Ianthella flabelliformis
Ianthella labyrinthus
Ianthella macula

Ianthella quadrangulata
Anomoianthella popae
Psammaplysilla purpurea
Pseudoceratina durissima

Fig. 6. Hypothetical relationships within the genus Ianthella
and within the Family Ianthellidae (Demospongiae:
Verongida). The hypothesised changes in character states
along each branch arc as follows. Characters are as in Table 1
and nodes have been given the reference numbers 1-5: nodes
l->2 (5b->a; 3d->a; le->b; 8b->a; 9b->a; 10b->a; 1 lb->a;
12d->c; 13d->c; 14e->c; 20b->a; 21a->b); nodes 2->3 (3a-
>c; 2b->a; 7b->a); nodes 3->4 (4b->a; 15d-> a); nodes 4->5
(lb->c; 12c->b; 13c->b; 14c->b; 16c->a; 17c->b); nodes 5-
>6 (6b->a); node l-> Pseudoceratina (17c->d); node 1-
>Psammaplysilla (16c->b); node 2->Anomoianthella (1b-
>d; 14c->d; 16c->d; 19a->b; 2Qa->c); node 4 ->I. reticulata
(15a->b); node 6 ->1. flabelliformis (3c->b; 16a->b); node 6-
>/. basta (lc->a; 12b->a; 13b->a; 14b->a; 15a->c; 17b->a;
18b->a).

172

Systematics and biogeography of the genus lanthella in the south-west Pacific

Figure 7. Zoogeographic distribution of lanthella and Anomoianthella (Family Ianthellidae) within the major Australian
marine faunal provinces (Bennetand Pope 1953; Hooper 1991), and a hypothesised evolutionary scenario for diversification
of the Ianthellidae.

The differentiation of species of lanthella and
their monophyly are well supported by morpho¬
logical data; characters that largely determine
the topology of the tree being the structure of the
skeleton, the nature of the fibres and density of
the soft tissues. The phylogenetic hypothesis
suggests that speciation of the genus lanthella
has proceeded from a three dimensional ances¬
tral sponge whose skeleton is a simple
anastomosing semi-dendritic network of very
large fibres, with dense collagenous tissue, to¬

wards a progressively more laterally compressed
body plan, with complex fibro-reticulation of
small regular meshes, ornamented surface
outgrowths, and diversity of pigmentation.

Models proposed for delineation of Austral¬
ian marine zoogeographic regions by Bennet
and Pope (1953), recognise five major divisions
which include two tropical provinces
(Dampierian and Solanderian), two warm-tem¬
perate provinces(Flindersian and Peronian) and
a cool-temperate province (Maugean) (Fig. 7).

Table 4. ZoogeographicaI distribution of Ianthellidae. Species distributions are listed within the major Australian marine
provinces (Bennet and Pope 1953) and other locations. The physical localities encompassed within these regions can be
found in Fig. 7. IND -Indian Ocean; SE ASIA-Southeast Asia, WCP-West Central Pacific.

IND

SE ASIA WCP

Dampierian

AUSTRALIA

Solanderian Flindersian Peronian

Anomoianthella popae
lanthella labyrinthus
lanthella flabelliformis
lanthella quadrangulata
lanthella reticulata
lanthella basta

x x(Guam) x

X X

X x

x(+south and north PNG)

173

P.R. Bergquist and M. Kelly-Borges

Recent interpretations of this model (Wilson
and Allen 1987; Hooper 1991) recognise broad
regions of overlap between zones as well as
narrow areas of endemism.

The known biogeographic distribution of
lanthella and Anomoianthella in the Australa¬
sian and Indo-West Pacific regions is given in
Table 4. Data were obtained from published
literature and from London’s Natural History
Museum, the Northern Territory Museum and
Australian Museum collections, McCauley et
al. (1992), and the authors’ unpublished collec¬
tions. In the absence of complete collections
from the faunal regions in question, a
biogeographic discussion can be given with
minimal confidence. However, the opportunity
exists to make some suggestions on speciation
events within the Ianthellidae, as the species in
this group have well defined and frequently
discrete distributions. Locality records have been
verified where possible by examination of mate¬
rial, and a reconstruction of the phylogeny of the
organisms is available.

The Dampierian region (mid-west to north¬
west Australian) has the greatest diversity of
Ianthellidae with two apparently endemic spe¬
cies, Anomoianthella popae and lanthella
labyrinthus (Table 4). From the Dampierian re¬
gion, /. basta extends north into Southeast Asia
and the West Central Pacific, and east across the
top of Australia into the Solanderian province
(Great Barrier Reef)- This is the most widely
distributed species of lanthella. Unlike I. basta,
the northern limit of/, flabelliformis is the south
coast of Papua New Guinea. It also extends east
and south into the Solanderian region, and may
occur further south in the cooler warm-temper¬
ate Peronian region (NSW coast), although as
mentioned earlier, this record is questionable.
lanthella quadrangulata is restricted to cool-
temperate southern waters of the Flindersian
province, and the Peronian province where it
overlaps in distribution with /. flab elliformis.
lanthella reticulata is restricted in distribution
to the Solanderian region but extends north onto
the south coast of Papua New Guinea with I.
flabelliformis, with a single record north of the
Papua New Guinea mainland.

These distributions suggest speciation and
dispersal of the genera Anomoianthella and
lanthella through an ancestor from the mid¬
west to north-western Australia (Dampierian
region), with /. basta and flabelliformis dis¬
persed into northern and eastern waters, and /.

quadrangulata into southern waters. The ab¬
sence of 1. reticulata from the Dampierian sug¬
gests that this species resulted later from a
northward dispersal event along the eastern
coast of Australia, through isolation from I.
quadrangulata by cooling southern waters. These
two species are very closely related morphologi¬
cally. This southward dispersal event has paral¬
lels in several microcionid genera (Hooper and
L6vi 1994).

The contention that the Dampierian region
might have been the centre of diversification of
lanthella and Anomoianthella is supported by
an exhaustive study of the Raspailiidae of Aus¬
tralia by Hooper (1991). Hooper found that the
fauna of the Dampierian province was extremely
diverse as compared with the fauna of the adja¬
cent Solanderian province, and the southern
Flindersian province,having many endemic spe¬
cies. L6vi (1979) also contends that the southern
Indonesian-northern Australian region has the
highest diversity forlndo-Pacific biogeographi-
cal provinces, and is the centre for dispersal for
Indo-Pacific sponges.

It is interesting to consider the hypothesised
phylogeny ofrelationships withinthe Ianthellidae
(Fig. 6) in the light of these zoogeographic
distributions. The phylogenetic reconstruction
also hypothesises that a common ancestor to
lanthella and Anomoianthella first led to the
derivation of both /. reticulata and /.
quadrangulata; which of these species came
first is equivocal in the analysis. A common
ancestor to the quadrangulata-reticulata group
gave rise to the endemic labyrinthus, and then to
the most recently derived and most widely dis¬
tributed basta and flabelliformis. Historical
events that might go towards explaining the
later speciation event are the warming of north¬
ern Australian waters during the Tertiary “jour¬
ney" northwards, and renewal of shallow-water
contact between Australia and Southeast Asia
during the Miocene.

FIELD KEY TO 1ANTHELLA

1 a. Symmetrical single or bilamellate fan, can
be stalked, up to 1 m high, 10-15 mm thick,
irregular rugose, knobbed surface, texture
harsh, compressible, yellow-green or blue.

. 2

b. Elongate fan or vase, occasional basal fanlets,
surface smooth to microconulose.3

174

Systematics and biogeography of the genus Ianthella in the south-west Pacific

c. Surface smooth and fleshy between well
spaced conules, sponge 5-10 mm thick,
fleshy, highly elastic and flexible, blue-
purple, yellow, orange and lilac, found on
the north-eastern Great Barrier Reef, and
predominantly in southern Papua New
Guinea. Ianthella reticulata

2 a. Surface covered in squat firm knobs and

very irregular ridges, sponge firm and
harsh to the touch, yellow-green or blue
with a whitish sheen, found in western,
northern, and eastern Australia and south¬
ern Papua New Guinea. Ianthella

flabelliformis

b. Surface outgrowths form concentric or laby¬
rinthine microconulose ridges, surface soft
and sponge pliable, stalked, found in west¬
ern Australia only Ianthella labyrinthus

3 a. Surface smooth , huge vertical rib-like

anastomosing fibres can be seen through
lamella, sponge fairly rigid in horizontal
plane, yellow, found in southern and west¬
ern Australian waters, and extends east¬
wards to Lord Howe Island. Ianthella

quadrangulata

b. Elongate fan or vase, occasionally with
basal fanlets, up to 1.5 m high, 1-3 mm
thick, very regular micro-conules forming
a ladder-like surface pattern, frequently
wavy, purple, green, yellow, orange or
blue, found throughout Australia, the In¬
dian Ocean and the Indo-West Pacific...
..... Ianthella basta

ACKNOWLEDGEMENTS

The authors thank the Motupore Island Re¬
search Department of the University of Papua
New Guinea, the Australian Institute of Marine
Science, and Walinde Resort, West New Brit¬
ain, for field support. For assistance with pho¬
tography, we thank Raewyn Eagar and Iain
MacDonald of the University of Auckland. Sup¬
port of the University of Auckland Research
Committee to Patricia R. Bergquist is acknowl¬
edged.

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Accepted 7 July, 1995

176

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:177-184

A NEW SPECIES OF LEPIDOTRIGLA (SCORPAENIFORMES: TRIGLIDAE)
FROM THE WATERS OFF NORTHERN AUSTRALIA.

LLUIS DEL CERRO AND DOMENEC LLORIS

Institut de Ciencies del Mar (C.S.I.C.),

Passeig Joan de Borbo, s/n 08039, Barcelona, Spain.

ABSTRACT

A new species of the family Triglidae, Lepidotrigla russelli sp. nov., from the waters
off the Northern Territory and Queensland (Australia) is presented. The new taxon is
compared with its similar relatives in the genus Lepidotrigla Gunther, 1860: L. faurei
Gilchrist and Thompson, 1914, and L. cadmani Regan, 1915. Brief comments on the
genus Lepidotrigla are included.

Keywords: Triglidae, Lepidotrigla russelli, sp. nov., taxonomy, new species, F.A.O.
Fishing Area 71, northern Australia, eastern Australia.

INTRODUCTION

As part of a general study on the systematics
of supra-specific taxa of the family Triglidae, a
large number of specimens collected in different
parts of die world have been examined. One par¬
cel of material examined contained a large
number of specimens coming from waters of
Indonesia and Australia. Among this material
was included an undescribed species clearly be¬
longing to the genus Lepidotrigla Gii nther, 1860.
The specimens used in the present study were
collected in Australian waters (F.A.O. Fishing
Area 71), mainly the Northern Territory (north
of Cape Wessel, north of Melville Island, north¬
east of Goulbum Island, off Jones Shoal, Bums
Shoal off Point Arrowsmith, east of van Diemen
Gulf) Western Australia (York Sound), and
Queensland (off Weipa). This new species, rep¬
resented by 17 specimens, is described below.

METHODOLOGY

The terminology of head spines mainly fol¬
lows Teague (1951), as well as Allis (1909) for
placement of spines on the cranial bones. The
terminology of the squamation follows Russell
et al. (1992).

Measurements and counts of body parts are a
combination of methods of Hubbs and Lagler
(1958), Teague (1951), Richards (1968), and
Richards and Saksena (1977), with the excep¬
tion of the following characters. The length of
the cleithral spine is measured from the poste¬
rior edge of the opercular flap to its rear end;
the lengths of the pectoral fin and its free rays
are measured from their respective superior
axils to their respective posterior tips; and the
length of the ventral fin is measured from the
ventral axil to the posterior end of the longest
ray.

None of the measurements and proportions
given here include the lengths of the rostral
projections which are measured separately. Their
length is measured from their tips to the pre¬
maxillary symphysis and the absolute values are
also given in Table 1. The head depth is meas¬
ured vertically from the posterior edge of the
orbit to the base of the head (preopercular bone).
The body depth is measured vertically, from in
front of the first anal ray and does not include the
crests at the bases of the dorsal fins.

Institution abbreviations used: Northern Ter¬
ritory Museum, Darwin, Australia (NTM) and
the Institut de Ciencies del Mar (C.S.I.C.), Bar¬
celona, Spain (IIPB), Museum National de
l’Histoire Naturelle, Paris (MNHN)).

177

L. Del Cerro and D. Lloris

Body measurement abbrevations are as fol¬
lows: TL, total length; SL, standard length; HL,
head length; OD, longitudinal diameter of orbit;
ML, maxillary length; CH, cheek height; and
10, interorbital distance.

SYSTEMATICS

Type material. HOLOTYPE-NTMS. 11953-
003, 194.1 TL (158.1 SL) mm, north-east of
Goulbumlsland (NorthemTerritory, Australia),
25 August 1986, depth 55 m.

PARATYPES - 16 specimens, size range
between 102.2 TL (84.8 SL) and 189.6 TL
(155.6 SL) mm: 1IPB 54/1993, 163.3 TL
(129.3SL) mm, 25th August, 1986, northeast of
Goulbum Island (NT), depth 55 m; IIPB 55/
1993, 186.9 TL (150.9 SL) mm, same data as
preceding; NTM S. 11953-012,178.9 TL( 144.9
SL) mm, same data as preceding; NTM S.
10053-002, 102.2 TL (84.8 SL) mm, October
1977, off van Diemen Gulf (NT); NTM S. 638,
107.1 TL (85.8 SL mm, July 1975, York Sound
(WA); NTM S. 1131,129.5 TL (115.5 SL) mm,
12 September 1975, off Jones Shoal (NT); IIPB
56/1993, 155.3 TL (120.3 SL) mm, off Cape
Wessel (NT); NTM S. 11614-007, 189.6 TL
(155.6 SL) mm, 11 May, 1985, same data as
preceding; NTM S.419, 155.2 TL (126.2 SL)
mm, 4 October 1975, Burns Shoal, off Pt
Arrowsmith (NT); NTM S.509,144.4TL (119.4
SL) mm, 18 May, 1983, N of Melville Island
(NT), 15 September 1975; NTM S.597, 165.8
TL (134.8 SL) mm, same data as preceding;
NTM S.598, 152.2 TL (123.2 SL) mm, same

data as preceding; NTMS.599,159.2 TL (130.2
SL) mm, same data as preceding; NTM S.600,
158.2 TL (126.2 SL) mm; NTM S. 578, 145.9
TL (117.9 SL) mm, September 1975, off Weipa
(Qld); NTM S.579, 144.4 TL (118.4SL) mm,
same data as preceding.

Other material. Other specimens used for
comparison were five syntypes of Lepidotrigla
laevispinnis Blache and Ducroz, 1960 (synony in¬
ked with Lepidotrigla cadmani Regan, 1915, by
Richards (1968: 81)), with the following cata¬
logue numbers: MNHN-1960-189 to MNHN-
1960-193, total and standard lengths range be¬
tween 212(172) and 190(158) mm, trawled west
of Pointe Noire (Congo Republic) by Blache and
Ducroz. Other capture data unknown.

Absolute measurements of the specimensstud-
ied are given in Table 1 for the new species. We
have not included proportional measurements
and percentages of the variables studied, leaving
this job to the reader.

Lepidotrigla Gunther, 1860
Lepidotrigla russelli sp. nov.

(Figs 1-4)

Diagnosis. Post-orbital groove (occipital or
post-frontal groove of some authors) incomplete,
reduced to form a short pit just behind the eyes.
Preopercular spine and keel are present. Length
of pectoral finsomewhatshorter thanhead length.
Body covered with firmly attached cycloid scales.
Throat, chest, breast and belly covered with small,
firmly attached cycloid scales.

Description. General counts and measurements
of holotype and paratypes are given in Table 1.

Fig. 1. Lateral view of the holotype NTM-S 11953-003 of Lepidotrigla russelli sp. nov..

178

Table 1. Absolute values of morphometric and meristic variables of the specimens of L. russelli sp. nov.. The meristic variable "1st. D1 Ray: Ser./Gran." indicates 0= smooth, S= serrated and
G= granulated.

A new gurnard from the waters off northern Australia

NTM-S-5 09

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179

L. Del Cerro and D. Lloris

Fig. 2. Lateral view of the head of the holotype NTM-S-11953-003 of Lepidotrigla russelli sp. nov.. Arrow marks the
preopercular spine and ridge.

Body robust anteriorly, slender posteriorly
and covered with large, firmly attached cycloid
scales, smaller on the chest, breast and belly,
numbering 12 to 13 between pelvic fin bases.
Lateral line scales 58 in holotype (57-63 in
paratypes), with their tubes slightly branched
(forming three branches); lateral line extending
onto the caudal fin forming a bifurcation. Both
sides of dorsal fin bases armoured with 22 erect
bony crests (22-23 in paratypes), flattened in
front and rising backwards, with eight erect
bony crests (seven to nine in paratypes, modally
eight) at the base of the first dorsal, and 14 at the
base of the second dorsal fin; some specimens
have one or two interdorsal crests.

Head rather large (3.1 times in SL), and
slightly spinulated. Post-orbital groove incom¬
plete, not crossing top of head, thus reduced to
a pit behind eyes. Nape scaled. Snout slightly
longer than orbit (2.7 in HL; 0.7 in OD), upper
profile scarcely concave. Rostral projection
much shorter than half length of orbit (27.3 in
HL; 7.2 in OD), with outer margin rounded
and armoured with several short spines. Orbit
rather large, somewhat longer than cheek depth
(3.8 in HL). Interorbital space deeply concave
(5.8 in HL) and much narrower than orbit (1.5
in OD). Maxillary not reaching to below ante¬
rior border of orbit (3.0 in HL). Teeth in both
jaws villiform; vomer edentate in holotype (two
paratypes with teeth) and palatines toothless.

Gill-rakers on first arch tubercle-like in shape,
well separated from each other and 8+2 rudi¬
ments in number. Pseudobranchials present.

First dorsal fin with nine spines (8-10 in
paratypes, modally 9), none reaching first ray of
second dorsal when depressed, with anterior
edge of first ray smooth (smooth or slightly
granulated in paratypes, but never serrated).
Second dorsal fin with 15 soft rays. Anal fin with
14 soft rays (13-15 in paratypes, modally 14),
inserted opposite origin of second dorsal fin.
Pectoral fin with 11 soft rays (10-11 in paratypes,
modally 11), moderate in length but always
shorter than head length, plus three free rays.
Pelvic fin well developed with one spine and five
soft rays, extending nearly to or slightly beyond
the vent (5.2 in SL). Caudal fin slightly
emarginate.

Spinulation. Cleithral spine stout and very
long; its length measured from the posterior
edge of opercular membrane to tip of spine is
slightly shorter than length of the orbit (4.1 in
HL) (3.3-5.0 in paratypes). Opercular spine
short. Preopercular spine and keel present and
conspicuous. Preocularspines absent in holotype
(0, 1 or 2 in paratypes), 1 postocular spine
present (1-2 in paratypes), sphenotic spine ab¬
sent (0-1 in paratypes), pterotic spine present,
parietal spine absent (0-1 in paratypes) and 1
nuchal spine present. Rostral, preorbital and
suborbital spines absent (sensu Teague 1951).

180

A new gurnard from the waters off northern Australia

Fig. 3. Dorsal viewofthe headoftheholotypeNTM-S-11953-003 of Lepidotrigla russelli sp.nov. Arrow marks the incomplete
post-orbital groove.

Fig. 4. Ventral viewofthe holotypc NTM-S-11953-003 of Lepidolrigla russelli sp.nov. showing the squamation of the chest,
breast and belly. Note the size of the scales.

Coloration in alcohol. Head and body yel¬
lowish brown. Body pale yellow in scaled areas,
but where the body scales are missing, the color
appears pinkish. Ventral region whitish. Pecto¬
ral fin black except for the two uppermost rays
which are pale, as are the free pectoral rays. In
some cases there are traces of scattered dark
spots on the spinous dorsal fin, spots which are
lacking in the holotype. Other fins pale.

Etymology. This species is named russelli
after Dr Barry C. Russell, Director of Research
and Collections at NTM.

DISCUSSION

The spinulation of the head and other charac¬
ters studied in the new species show a certain
degree of variability as indicated above in the
description. Nevertheless, all these data have
been included here in order to follow the stand¬
ardized descriptions of the family given by other
authors.

Two specimens, NTM S. 11614-007 and NTM
S.579, have teeth on the vomer in the sense that
this bone had these teeth formed by very small

181

L. Del Cerro and D. Lloris

spinulations, whilst specimens of different spe¬
cies the genus Lepidotrigla had papillae on the
dermal tissue covering the bone, with the latter
being smooth. This could have confused some
authors who identified these papillae as true
vomerine teeth. Thus, in our opinion, the pres¬
ence or absence of teeth on vomer, as given in
some works on the family, should be read with
some caution.

Due to the presence of an incomplete post¬
orbital groove and pectoral fin length being less
than the length of the head, this species resem¬
bles the following species, including those con¬
sidered as valid by Richards (1992: 54-55, Table
5): L. abyssalis Jordan and Starks, 1904; L. a lata
(Houttuyn, 1782); L. alcocki Regan, 1904; L.
cadmani Regan, 1915; L. faurei Gilchrist and
Thompson, 1914;L.g«e«//im'Hilgendorf, 1879;
L. longifaciata Yatou, 1981; L. tnarisincnsis
Fowler, \93Z\L.modesta Waite, 1899 \L.oglina
Fowler, 1938;L. spiloptera Gunther, 1880; L.spin-
asaGomon, 1987 and L. umbrosa Ogilby, 1910.

The presence of scales on the throat, chest,
breast and belly separates this species from all
those in the list above except for L. alata, L.
faurei and L. cadmani. The former species has
the rostral appendages formed into a single,
long, stout spine, which some authors have used
to include L. alata in the genus Pachytrigla
Fowler, 1938, and is thus easily separated from
the new species. Lepidotrigla faurei has been

Fig. 5. Detail in dorsal view of the predorsal area of
Lepidotrigla russelli sp. nov. (A) and L. cadmani (B).

confused due to the distribution of the scales on
the throat, chest, breast and belly. There is no
reference to this character in the original de¬
scription given by Gilchrist and Thompson
(1914: 75). Smith (1934: 322, 324-325) de¬
scribes the species L. faurei as having scales on
the breast and differentiates this species from L.
natalensis Gilchrist and Thompson, 1914, by
the breast squamation. The same procedure was
followed by Richards (1968: 81) in his descrip¬
tion of the new species L. carolae, separating
this species from L. cadmani by the differences
in breast squamation. Blache and Ducroz( 1960:
208), in their description of L. laevispinnis (= L.
cadmani after Richards (1968: 81) compare
their new species with L. faurei, stating that both
species have scales on the breast and in the inter-
ventral area. Richards (1992: 54, Table 5), used
as a reference work for our list of valid species,
writes “naked” in the column “breast
squamation” for the species L. faurei (where he
considers L. natalensis Gilchrist and Thompson,
1914, and L. stigmapteron Fowler, 1934, as
synonyms of the nominal). Besides our interest
in maintaining Richards (1992) as the only
reference list of valid species of the genus
Lepidotrigla, we cannot ignore clear references
to breast squamation of the above mentioned
species in previous works. Taking into account
that Richards (1992: 54, Table 5) had no access
to type material of L. faurei, we have not consid¬
ered the data on squamation given by this author
for that species.

Thus, Lepidotrigla russelli sp. nov. differs
from L. faurei because the latter shows the
following characters: the first three dorsal rays
have serrations in their anterior edges (smooth
or granulated in the new species); there is no
spine or keel present on the preopercular bone
(both are present in the new species); and the
body scales are ctenoid but the ventrals are
cycloid (scales always cycloid in the new spe¬
cies).

Lepidotrigla cadmani is very similar to the
new species, both species having biometric char¬
acters which are very close, except for the cheek
height and the interorbital distance versus the
maxillary length. In L. cadmani, the height of
the cheek is always greater than the maxillary
length (ML= 0.8 to 0.9 in CH for the syntypes of
L. laevispinnis) but much smaller than the max¬
illary length in the new species (ML= 1.4 to 2.8
in CH). The interorbital distance is greater in L.
cadmani (ML= 1.3 to 1.5 in IO for the syntypes

182

A new gurnard from the waters off northern Australia

of L. laevispinnis) than in the new species (ML=
1.6 to 2.1 in IO) There is clearly no overlap
among the ranges of both species for the first
character mentioned but there may be a very
slight one in the second.

From the morphological and meristic point of
view, both species are easily separated.
Lepidotrigla cadmani has no preopercular keel
(present in the new species), has an inconspicu¬
ous preopercular spine (distinct preopercular
spine), semi-circular predorsal area (predorsal
area pinched-in, with a narrow anterior exten¬
sion, (Fig. 5)), five tubules in the scales of the
lateral line (three tubules in the scales of the
lateral line), body scales ctenoid, cycloid on
breast (all scales cycloid), and four to five scales
between pelvic fins (12 to 13 scales between
pelvic fins). While reviewing literature on the
Triglidae of the area, we found a particularly
interesting species cited by Goerfelt-Tarp and
Kailola (1983: 117) as Lepidotrigla sp. 4. Un¬
fortunately, the description given by these au¬
thors is too short and incomplete to assign this
or these specimens to any known species, though
the meristic characters agree with those of the
new species, as do the unusual presence of scales
on the breast. These features may indicate that
the new species described in this paper and
Lepidotrigla sp. 4 could very well be the same.

Some authors give particular importance to
the shape of the spines of the preorbital bones.
We have been revising the genera of the family
Triglidae (including Peristediidae) since 1990,
and in our opinion, not too much importance
should be given to this feature, though it is
helpful as complementary information for dif¬
ferentiation of some groups. This may also be
true for coloration of the first dorsal fin and the
presence or absence of teeth on the vomer. We
have found variation in the first feature in many
specimens of different species of this family. As
has been indicated in the description of the new
species, the pattern of spots on the first dorsal is
not constant. The same occurs with the vomerine
teeth, as their presence is variable, as has been
stated by several authors such as Smith (1934:
323,327) for the genus Lepidotrigla , and Hubbs
(1959: 314) for the genus Pterygotrigla Waite,
1899, among others.

Yato (1981:263) states: ‘ Identification of the
gurnards referable to the genus Lepidotrigla is
extremely difficult because of the taxonomic
characters vary strikingly with growth stage and
individuality’. The observation perhaps also

applies because the original descriptions and
those given by subsequent authors often give
differing information on the species treated.
Such varying information is often difficult to use
when making comparisons, or attempting an
accurate species determination, as in some cases
the characters given in various papers differ
widely and are impossible to compare. We also
believe that to revise all genera of the family is
necessary; works of wider scope are likely to
provide a better understanding of the taxonomy
of the Triglidae sensu lato.

ACKNOWLEDGEMENTS

We are indebted to Dr Barry C. Russell, of the
Museum and Art Gallery of the Northern Terri¬
tory, Darwin, who kindly loaned a large amount
of triglid material to twohitherto unknown men,
extended the period of loan when necessary and
did his best to attend to our requests.

We wish also to give here our grateful thanks
to Drs Bernard Seret and Guy Duhamel from the
Museum National d’Histoire Naturelle, Paris,
who, besides many other favours, rapidly re¬
sponded to our request for triglid material for
comparison with the rare species we found in the
Indo-Australian material.

We are grateful to Ms Barbara Brocklehurst
who revised the final English version and also to
Ms Helen K. Larson, editor of The Beagle, and
Dr W.J. Richards, for their comments on the
manuscript.

REFERENCES

Allis, E. P. 1909. The cranial anatomy of the mail¬
cheeked fishes. Zoologica, Stuttgart 57: 1-219.

Blache, J. and Ducroz, J. I960. Lepidotrigla
laevispinnis n. sp. (Pisces, Perciformi,
Scorpaenoidei, Triglidae), poisson nouveau du
plateau continental congolais. Bulletin du
Museum National d’Histoire Naturelle, 2e. Serie
32(3): 205-208.

Fowler, H.W. 1934. Fishes obtained by Mr. H.W.
Bell-Marley chiefly in Natal and Zululand in
1929 to 1932. Proceedings of the Academy of
Natural Sciences of Philadelphia 86: 405-514.

Fowler, H.W. 1938. Descriptions of new fishes ob¬
tained by the United States Bureau of Fisheries
Steamer “ Albatross ” chiefly in Philippine Seas
and adjacent waters. Proceedings of the United
States National Museum 85(3032): 31-135.

183

L. Del Cerro and D. Lloris

Gilchrist, J.D.F. and Thompson, W.W. 1914. De¬
scription of fishes from the coast of Natal. (Part
IV). Annals of the South African Museum 13(3):
65-95.

Gloerfelt-Tarp. T. and Kailola, PJ. 1984. Trawled
fishes of southern Indonesia and north Western
Australia. Australian Development Assistance
Bureau: Canberra.

Gomon, M.F. 1987. New Australian fishes. Part. 6.
New species of Lepidotrigla (Triglidae),
Choerodon (Labridae) and Zebrias (Soleidae).
Memoirs of the Museum of Victoria 48( 1): 17-
23.

Gunther, A. 1860. Catalogue of the Acanthopterygian
-fishes in the collection of the British Museum.
2. London. XXI, 548 pp.

Gunther, A. 1880. Report on the shore fishes pro¬
cured during the voyage of H.M.S. Challenger in
the years 1873 - 1876. Challenger Report, Zool¬
ogy 1(6): 1-82.

Hilgendorf, F.M. 1879. Diagnosen neuer Fischarten
von Japan. Sitzungsberichte der Gesellschaft
Naturforschender Freunde zu Berlin: 105-111.

Houttuyn, M. 1782. Beschryving van eenige Japa¬
nese visschen en andere Zee-Schepzelen.
Verhandelingen uitgegeeven door de Hollandse
Maatschappye der Wetenschappen te Haarlem
20(2): 311-350.

Hubbs, C.L. 1959. Initial discoveries of fish faunas
on seamounts and offshore banks in the eastern
Pacific. Pacific Science 13: 311-316.

Hubbs, C.L. and Lagler, K.F. 1958. Anatomical
features and terms and methods of counting and
measuring. In: Fishes of the Great Lakes Region.
Bulletin of the Cranbrook Institute of Science
26: 19-26.

Jordan, D.S. and Starks, E.C. 1904. List of fishes
dredged by the steamer ‘•Albatross’’ off the coast
of Japan in the summer of 1900, with descrip¬
tions of new species and a review of the Japanese
Macrouridae. Bulletin of the United States Com¬
mission for Fisheries for 1902 22: 577-630.

Ogilby, J.D. 1910. On some new fishes from the
Queensland coast. Proceedings of the Royal
Society of Queensland 23: 85-139.

Regan, C.T. 1904. Descriptions of three new marine
fishes from South Africa. Annals and Magazine
of Natural History 7 (14): 128-130.

Regan, C.T. 1915. A collection of fishes from Lagos.
A nnals and Magazine of Natural History 8(15)-
124-130.

Richards, W.J. 1968. Eastern Atlantic Triglidae (Pis¬
ces, Scorpaeniformes). Atlantide Report 10: 77-
114.

Richards, W.J. 1992. Comments on the genus
Lepidotrigla (Pisces, Triglidae) with descrip¬
tions of two new species from the Indian
OcemMulletin of Marine Science 51(1): 45-65.

Richards, W.J. and Sakena, V.P. 1977. Systematics
of the gurnards, genus Lepidotrigla (Pisces,
Triglida), from the Indian Ocean. Bulletin of
Marine Science 27 (2): 208-222.

Russell, M., Grace, M. and Gutherz, E.J. 1992. Field
guide to the searobins ( Prionotus and Bellator)
in the western north Atlantic. National Oceanic
and Atmospheric Administration Technical Re¬
port NMFS Circular 107.

Smith, J.L.B. 1934. The Triglidae of South Africa.
Transactions of the Royal Society of South Af¬
rica 22(4): 321-336.

Teague, G.W. 1951. The sea-robins of America. A
revision of the triglid fishes of the genus
Prionotus. Communicaciones Zooldgicas del
Museo de Historia Natural de Montevideo 3(61)'
1-59.

Waite, E.R. 1899. Scientific results of the trawling
Expedition of H.M.C.C.S. “Thetis". Fishes.
Memoirs of the Australian Museum 4: 1-132.

Yatou, T. 1981. A new triglid fish, Lepidotrigla
longifaciata, from Japan. Japanese Journal of
Ichthyology 28(3): 263-266.

Accepted 20 January, 1994

184

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:185-192

A NEW FISH PARASITE (COPEPODA: SIPHONOSTOMATOIDA:
CALIGIDAE) FROM THE TIMOR SEA, AUSTRALIA.

KUNIHIKO IZAWA
Faculty of Bioresources, Mie University,

1515 Kamihama, Tsu, Mie 514, Japan.

ABSTRACT

A new species of parasitic copepod, Pseudopetalus timorensis n. sp., is described based
on a single specimen (female) obtained from a pelagic fish, possibly a flying fish
(Exocoetidae) or a long tom (Belonidae), collected by a dip net at night in the Cartier
Reef, Timor Sea, Australia, in May, 1992. The species is distinguishable from its
congeners in having a proportionally long aliform abdomen with strongly curled edges.

Keywords: new species, fish parasite, Copepoda, Siphonostomatoida, Caligidae,
Pseudopetalus, Timor Sea, Australia.

INTRODUCTION

A parasitic copepod of the genus
Pseudopetalus Pillai, 1962 (Caligidae), depos¬
ited in the Museum and Art Gallery of the North¬
ern Territory, Darwin, is here described as a new
species. Currently fourspecies of Pseudopetalus
are known (Table 1). They are parasitic in the
buccobranchial cavities ofclupeid and belonid
fishes in Indo-Pacific waters. This is the first
record of Pseudopetalus from Australian waters.

Abbreviations used in the figures are: A’,
antennule; A”, antenna; Lu, lunule; Mt, mouth
tube; Mx’, maxillule; Mx”, maxilla; Mxp,
maxilliped; Pap, postantennal process.

MATERIAL AND METHODS

The specimen was found loose in a mixed
collection obtained by using a dip net. The col¬
lection was made at night at the west end of
Cartier Reef, Timor Sea, Australia (12°32.2’S
123°31.7 ’E) by Dr B. C. Russell on 6 May 1992.
The host fish is unverifiable, but it was suspected
to be either a flying fish (Exocoetidae) or a long
tom (Belonidae). The specimen, preserved in
alcohol, was examined in lactic acid under both
dissecting and compound microscopes. Draw¬
ings were made using a drawing tube.

SYSTEMATICS

Caligidae

Pseudopetal us Pil lai
Pseudopetalus timorensis n. sp.

(Figs 1-4)

Type material. The holotype is deposited in
the Museum and Art Gallery of the Northern
Territory, Darwin (NTM Cr. 009414).

Description. Female: Total length (excluding
egg sacs) 8.9 mm, greatest width about 2.2 mm.
Body (Fig. 1) elongated, about four times longer
than wide, divisible into three parts: anterior oval
cephalothorax; median enlarged, flexible geni¬
tal complex, which attaches to cephalothorax
through the small fourth pediger; and posterior
prominent troughlike abdomen, in which egg
sacs are held. Egg sac nearly straight, 5.5 mm
long and 0.4 mm in diameter, containing about
67 flattened eggs arranged uniserially.

Cephalothorax longer than wide (1.7 X 1.3
mm), with brownish pigment spots scattered
over convex dorsal shield with relatively narrow
membranous flange along border accompanied
sparsely by sensory hairs (Fig. 2 A). Four areas of
dorsal shield well demarcated by sutures; ce¬
phalic area extending about two thirds of
cephalothoracic length, with frontal plate carry¬
ing pair of lunules. Frontal plate indented at

185

K. Izawa

middle of anterior margin, with ring-like sclerite
at middle of ventral side and membranous flange
on anterior margin between lunules. Lunules
consist of rugose adhesive disc and marginal
membrane (Fig. 2A). Thoracic area wider than
long, with posterior sinuses and gently curved

posterior margin. Lateral areas small, bearing
sensory crypt posterolaterally.

Fourth pedigerous somite small, wider than
long (0.2 X 0.4 mm) and tapered anteriorly.

Genital complex 3.2 mm long (including pos-
teriorlobes) and 1.6mmwide(inposteriorthird).

Fig. 1. Pseudopetalus timorensis n. sp„ female. A, habitus, dorsal; B, same, ventral; C, same, lateral.

186

New parasitic copepod from northern Australia

Anterior eighth of genital complex narrow and
cylindrical and succeeded by slightly broadened
portion with winglike lateral expansions. Poste¬
rior two-thirds gradually broadened posteriorly
and ending in pair of lamelliform lobes, at their
bases, a pair of genital processes bearing the
genital pore medially. No rudiments of fifth and
sixth legs on genital complex.

Abdomen large, 4.5 mm in length and com¬
posed of two segments; first segment long,
broadly expanded laterally and strongly curled
ventrally, with lateral margins nearly touching
each other and posterior margin extending be¬
yond small second segment. Latter shorter than
wide, carrying caudal rami. Abdomen attached
to posterodorsal part of genital complex above
genital processes. Ventral side of abdomen
slightly inflated along midline; gut visible by its
brownish contents.

Antennule (Figs 2A, 2B) two-segmented, 0.3
mm long. First segment somewhat flattened
dorsoventrally and broadened distally, with 26
plumose setae, of which 19 are arranged in two
rows ventrolaterally, six dorsolaterally and one
mediodistally. Second segment rodlike, with 12
simple setae and one aesthetasc distally and one
seta medially.

Antenna (Fig. 2A) four-segmented. First seg¬
ment arising from sternal swelling and laid in
sternal groove, unarmed. Second segment short,
with spiniform process projected
posteromedially. Third segment robust, unarmed;
and fourth segment bearing strong claw with two
setules, one ventral near base and another mid¬
way on segment.

Hooked postantennal process (Fig. 2A) later¬
ally to antennal base, with three basal setiferous
papillae.

Mouth tube (Fig. 2A) 0.45 mm long and 0.08
mm wide, reaching far beyond basesof maxillae,
with membranes around opening.

Mandibles originating laterally to mouth tube
and inserting their distal portions laterally into
mouth tube.

Maxillules (Fig. 2A) flanking mouth tube and
mandibles, composed of hook and papilla
tipped with three setules.

Maxilla (Fig. 2A) slender, three-segmented:
first and second segments naked; third segment
slender, with flabellum on distal fourth of ante¬
rior side and two subecjual, feeble membra¬
nous claws at tip.

Maxilliped (Fig. 2A, Mxp; 2D, 2E) stout,
three-segmented: first segment (corpus

maxillipedis) robust, with pointed process and
ridgelike bulge on medial side, and small swell¬
ing on posterior side; distal two segments form¬
ing strong claw, with small knob and setule at
midlength.

Sternal furca (Fig. 2A, C) between bases of
maxillipeds, with somewhat elongate base and
moderately diverging fork, without flange.

First leg (Fig. 3A) smaller than following
pairs, composed of two-segmented protopod and
exopod. Coxa with setule laterally on anterior
side; basis with digitiform rudiment of endopod
on unsclerotized distal portion, long, plumose
lateral seta, short plumose seta on mediodistal
corner, and spinules scattered on anterior side
mediodistally. First exopod segment elongate,
with small subterminal, lateral spine and row of
spinules on medial edge; second segment with
four pinnate setae and three distal spines; outer¬
most spine with membranes and other two spines
with accessory blades.

Second leg (Fig. 3B) large and elaborate.
Intercoxal plate flaplike, two times as wide as
long, with broad membranous flange along free
margin. Protopod two-segmented: coxa broad
but short, with membrane along lateral free mar¬
gin and medial plumose seta and unarmed setule;
basis narrower, but longer than coxa, with sim¬
ple lateral seta, broad membrane and fine un¬
armed seta on distal free margin. Exopod
three-segmented: first segment nearly as long as
distal two segments combined, with spine ac¬
companied by flabellum at base on distolateral
swelling, pinnate seta mediodistally, and mem¬
branous flange and row of setules on outer and
medial margins, respectively; second segment
short, with spine on outer distal swelling and
pinnate seta medially; and third segment with
three spines and five pinnate setae. All spines of
exopod with membranes, but distalmost spine
with pinnules on medial side. Endopod three-
segmented; first segment with row of spinules on
outer distal margin and medial pinnate seta; second
segment largest, armed with 14 stout cuspid-like
processes arranged in two rows along outer mar¬
gin and two medial pinnate setae; third segment
small and semicircular, with six pinnate setae.

Third leg (Fig. 4A) consists of broad protopod
and small rami. Apron longer than wide, with
membranous flange along free margin and
digitiform sclerotized lateral process. Coxa and
basis almost completely fused together, armed
with medial spinous pad on ventral side, spinulose
patches on ventrolateral side, fine sculpture on

187

K. Izawa

proximal dorsolateral margin, broad membra¬
nous flange on lateral margin, plumose lateral
seta on distal comer, pinnate seta on mediodistal

comer, and membranous flange accompanied by
two setules on mediodistal free margin. Exopod
three-segmented: first segment small, with sin-

188

New parasitic copepod from northern Australia

Fig. 3. Pseudopetalus timorensis n. sp„ female. A, first leg, anterior, B, second leg, anterior.

gle spine (missing in Figure 4A); second seg¬
ment with row of fine setules and simple, weak
outer spine, and pinnate medial seta; third seg¬
ment with row of setules on outer edge, three
spines, and four pinnate setae. Endopod
two-segmented: first segment flaplike, broadly
articulated to basis, with row of setules along
entire margin and pinnate medial seta; second

segment with row of setules laterally and six
pinnate setae.

Formulae of spines and setae of legs 1 -3 as
follows:

Leg 1 Coxa 0-0 Basis 1-1 Exopod 1-0; 111,1,3

Leg 2 Coxa 0-1 Basis 1-0 Exopod 1-1; 1-1; 111,5

Endopod 0-1; 0-2; 6
Leg 3 Coxa+Basis 1-1 Exopod 1-1; 1-1; 111,4

Endopod 0-1; 6

189

K. Izawa

Fig. 4. Pseudopetalus timorensis n. sp., female. A, third leg, ventral; B, fourth leg, anterior; C. posterior portion of
abdomen and a caudal ramus, ventral.

Fourth leg (Fig. 4B) three-segmented,
uniramous; first segment (sympod) as long as
distal two segments combined, with plumose
distolateral seta; second segment with distal spine
and fine setule on outside; third segment with

four spines; all spines clawlike, with fine mem¬
branes and accompanied by flabellum at base.

Caudal ramus (Fig. 4C) rectangular, tipped
with three long pinnate setae and three small
setae.

190

New parasitic copepod from northern Australia

DISCUSSION

Pseudopetalus timorensis, n. sp., is easily dis¬
tinguished from its congeners by the proportions
and appearance of the abdomen. As indicated in
Table 1, the new species has the greatest propor¬
tion of abdomen, comprising more than half of
its total body length. In all other species of
Pseudopetalus, the abdomen does not curl as in
the new species, although it was shown gently
curved ventrally in P. caudatus (Gnanamuthu,
1950, Fig. 1). The abdomen of P. formicoides
(Redkar, Rangnekar and Murti, 1949) and P.
dussumieri (Rangnekar, 1957) is more slender
than that of P. caudatus and P. denticulatus
(Shen, 1957). In P. caudatus and P. timorensisn.
sp., the aliform posterior edge of the first seg¬
ment of the abdomen reaches far beyond the
second abdominal segment, but no such forma¬
tion is found in P. formicoides, P. denticulatus,
and P. dussumieri. Though similar to P. caudatus
among its congeners in having a broad abdomen
with posterior lobes, P. timorensis n. sp. differs
from P. caudatus (Pillai 1968,1985) in armature
of the appendages as follows: in P. timorensis n.
sp. the maxilliped has a pointed tooth on myxa
other than a ridgelike bulge instead of two teeth.
The antennal posterior process projects
posteromedially in P. timorensis n. sp., while it
projects posterolaterally in P. caudatus. The first
and second endopod segments of leg 2 are armed
with a row of fine spinules and 14 teeth on the
lateral margins respectively in P. timorensis n.
sp., but in P. caudatus these segments are armed

with three to four teeth and 10 to 12 teeth respec¬
tively. The teeth of the second segment of the
former are thicker than those of the latter and
these teeth arise contiguously at the bases in¬
stead of at regular intervals. Moreover, in P.
timorensis n. sp. three medial setae of the distal
exopod segment of leg 1 are pinnate and longer
than distal spines, but these setae are replaced by
short simple setae in P. caudatus. The setae on
the terminal segment of leg 1 show various
degrees of reduction in this genus. They are
rudimentary in P. formicoides (Redkar et al.

1949; Pillai 1962) and P. caudatus (Gnanamuthu
1950; Pillai 1962), about as long as the segment
in P. denticulatus (Shen 1957) and P. timorensis
n. sp., and longer than the segment in P.
dussumieri (Rangnekar 1957; Pillai 1968).

P. denticulatus was treated as a variety of P.
formicoides by Pillai (1962) and then as a syno¬
nym of the latter by Pillai (1985). However, it
must be considered a valid species, as it is dis¬
tinct from the latter in the shape of the abdomen,
armature of the maxilliped myxa, relative length
of setae on the terminal segment of leg 1, and
shape and number of cuspid-like processes of the
second endopod segment of leg 2.

Males are unknown for this genus.

ACKNOWLEDGMENTS

The author expresses his thanks to Drs A. J.
Bruce and B. C. Russell of the Museum and Art
Gallery of the Northern Territory, Darwin, for
giving him an opportunity to examine the speci-

Table 1. Species of Pseudopetalus, with morphometric comparisons. BL, total body length; GL and GW, length and
width of genital segment; AL and AW, length and width of abdomen.

Taxon

Reference

Host

GL/BL

GW/GL

AL/BL AW/AL

(mm)

P. formicoides

Redkar et al. (1949)

on Dussumieria acuta
from Bombay, India Ocean

9.0

0.40

0.45

0.47

0.49

P. formicoides

Pillai (1962)

on Sardinella fimbriata,
from Madras, Indian Ocean

10.2

0.41

0.35

0.49

0.38

P. caudatus

Gnanamuthu (1950)

on Dussumieria acuta
from Madras

7.4

0.44

0.52

0.46

0.81

P. caudatus

Pillai (1962)

on Dussumieria hasseltii
from Trivandrum, Indian Ocean

8.5

0.39

0.52

0.44

0.80

on Ablennes hians from
Trivandrum

9.1

0.44

0.42

0.95

P. denticulatus

Shen (1957)

on Dussumieria hasselti from
Hai-Nan Is., South China Sea

6.7

0.38

0.55

0.50

0.64

P. dussumieri

Gnanamuthu (1957)

on Dussumieria acuta from
Bombay

6.2

0.31

0.73

0.43

0.28

P. dussumieri

Pillai (1968)

on Dussumieria hasselti from
Trivandrum

5.2

0.37

0.68

0.39

0.52

P. timorensis n.sp.

this paper

on flying fish or long tom,

Cartier Reef, Timor Sea

8.9

0.37

0.48

0.52

(0.47)

191

K. Izawa

men. Thanks are also due to Drs J.-S. Ho of the
California State University, Long Beach, andZ.
Kabata, Pacific Biological Station, Nanaimo, for
their critical reading of and comments on the
manuscript.

REFERENCES

Gnanamuthu, C. P. 1950. Parapetalus caudatus n.
sp., a cope pod parasitic on Dussumieria acuta,
from Madras. The Proceedings of the Indian
Academy of Sciences 31(2), Sec. B: 125-133.

Pi llai, N. K. 1962. A revi sion of the genera Parapetalus
Steenstrup & Liitken and Pseudopetalus nov.
Crustaceana 3(4): 285-303.

Pillai, N. K. 1968. Additions to the copepod parasites
of South Indian fishes. Parasitology 58: 9-36.

Pillai, N. K. 1985. Fauna of India: Parasitic copepods
of marine fishes. Zoological Survey of India:
Calcutta.

Rangnekar, M. P. 1957. Caligus dasyaticus sp. nov.
and Caligus dussumieri sp. nov., (Copepoda)
parasitic on Bombay fishes. Journal of the Uni¬
versity of Bombay 25(5), Sec. B: 16-22.

Redkar, M., Rangnekar, P. G. and Murti, N. N„ 1949.
Four new species of parasitic copepods from the
marine fishes of Bombay. Journal of the Univer¬
sity of Bombay 18(3), Sec. B: 36-50.

Shen, C.-J. 1957. Parasitic copepods from fishes of
China part II. Caligoida, Caligidae (1). Acta
Zoologica Sinica 9(4): 351-377, Plates I-XI.

Accepted 24 April, 1995

192

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:193-208

WHO ARE THE BAY IN I ?

IAN MCINTOSH

School of Social Sciences, Northern Territory University,
Darwin, NT 0909, Australia.

ABSTRACT

In the literature, the first non-Aboriginal visitors to the Arnhem Land coast (pre -
Macassan) are referred to as Bayini. The stories of the Bayini have meaning only when
seen through the mythology of the ancestral being Birrinydji, who, in a “cargo” type
perspective, is the foundation of the wealth of non-Aborigines. At the "beginning of
time", Aborigines possessed this wealth, but through misadventure this was lost, and
now “whites” control the production and distribution of material goods. The mythol¬
ogy of Birrinydji is constituted in the ever-changing nature of relations between
Aborigines and non-Aborigines. In an examination of a Rag treaty proposal by
members of the Warramiri clan of Elcho Island, it is apparent that accounts of the past,
seen through this body of law, are constandy changing. This fact, combined with the
almost innumerable possible variations in accounts emphasising either “inside” or
“outside” myth or history or combinations thereof, suggests that while a complex past
is being evoked in Bayini narratives, great caution is needed when attempting to make
definitive statements about it.

Keywords: Aboriginal art. Aboriginal history. Aboriginal myth, Australian flag,
Bayini, Birrinydji, cargo cult, identity, Macassan, northern Australia, reconciliation,
representation, the “Other", treaty.

INTRODUCTION

Mystery surrounds the identity of the Bayini,
a group of people that Yolngu (Aborigines from
north-east Arnhem Land) regard as the forerun¬
ners of Macassans. By “Macassan”, I am refer¬
ring to those peoples who were engaged in the
trepang industry on the northern Australian
coast from at least 1700, and whose historical
existence is not in question. Aborigines clearly
differentiate the Bayini from the Macassans,
and herein lies the problem from an anthropo¬
logical viewpoint. Aboriginal accounts locate
the “historical” Bayini in Arnhem Land, but the
stories are also connected with Macassar or
Ujung Pandang, and the trepang industry (Bemdt
and Bemdt 1954:37; Mountford 1956-64:334).
As the historian Macknight (1972: 313) indi¬
cates, much of the terminology associated with
them is also Macassan.

Despite this, speculation on the identity of the
Bayini is rife. Some writers say they were
Macassan, Sama-Bajau or sea nomads (Bemdt
1965:5), while others suggest they were

Gudjeratis from India (Halls 1965: 4), Chinese
(Levathes 1994; Worsley 1955: 2), Dutch, or
even Portuguese (Mountford 1956-64: 334). In
one extreme case, they were considered to be
voyagers from the moon and were brought to the
attention of the science fiction writer Von
Daniken (E. Saffi pers. comm. 1993). It is fair to
say that this question represents one of the
longer standing puzzles of northern Australian
history and anthropology (Capell 1965: 68).

In seeking to contribute to an answer to this
puzzle, I compare and contrast ten Aboriginal
art works pertaining to external influences and
relate these to ever-changing nature of accounts
of “totem” hunters, Bayini and Macassans in the
literature.

Macknight (1972: 317; 1986: 72) called for
an analysis of the way the memory of the trepang
industry has been transformed and put to use in
contemporary Aboriginal politics. I suggest that
the Bayini are in the image of how some Abo¬
riginal groups are believed to have once existed
at the “beginning of time” (in the Wangarr or
“Dreaming”). In an almost archetypal “cargo

193

I. McIntosh

cult” scenario (e.g. Burridge 1971), Aborigines
once possessed the wealth and technologies of
non-Aborigines. Following contact with
Macassans, poor“black” Aborigines found them¬
selves to be dominated by rich “whites”.

I contend that the actions of Macassans and
possibly other voyagers to the Australian coast
have become the basis of a sacred law relevant in
contemporary Aboriginal dealings with non-
Aborigines. From this perspective, I suggest that
the search for a single group by the name Bay ini,
is misplaced. More important is it to ask, why do
some people use this category, while others do
not? The mythology and associated historical
descriptions of the Bayini are an example of
Aboriginal imagination at work in a continuing
construction of identity in the context of an ever-
changing relationship with the “other”'. The
term Bayini is an “outside” or public label 2 ,
relevant in particular contexts in discussions
about such relationships.

HISTORICAL OVERVIEW

The most authoritative accounts of the
Macassan era are Macknight’s (1976) The voy¬
age to Marege, and the work of R.M. and C.H.
Bemdt (1954) in Arnhem Land, its history and
its people. While Macknight looked specifically
at the history and organisation of the trepang
industry, the Bemdts (1954: 64) attempted to
show that there were a numberof discrete phases
of external contacts. According to informants in
the 1940s, Aborigines were subjected to four
separate waves of visitation prior to the arrival of
Japanese pearlers and Europeans this century.

Bemdt and Bemdt (1954:64) suggest that for
countless centuries, Aborigines had been aware
of a people known as Wurramala. Song cycles
concerning their activities had their origins in
the tidal flotsam from the islands to the north,
and also by irregular visits by peoples from there
(Bemdt 1948: 103). Accounts are ambiguous. In
some interpretations, these "totem" hunters are
associated with the “land of the dead” and assist
in the passage of the soul of deceased Aborigines
to this place. In others, the songs are said to be
about reflections on travel aboard Macassan

praus (Bemdt 1948). Informants were divided
on whether eastern Indonesia and/or Torres
Strait was involved (Bemdt 1978/79: 65-67;
McIntosh 1995).

Stories of the exploits of “totem” hunters are
associated with what Morphy (1990) referred to
as myths of creation. Representations in art are
few and more often than not, the presence of the
“other” is inferred in paintings depicting the

Fig. I. The body of the whale captured by the “totem" hunter
Wurramala. north of Elcho Island. The whale’s backbone is
depicted on the left hand side. By Bapayili of the Wolkara
clan. Whether the stories associated with these “totem" hunt¬
ers reflect early Macassan encounters or a separate wave of
historical contact, is unknown. Courtesy of tire Museum and
Art Gallery of the Northern Territory (Abart366).

1 The constitution of the "other” by anthropologists and by indigenous peoples is a major issue in contemporary anthropology
(Fabian 1983). This paper discusses the encounter of each with the other, i.c. Aboriginal representations of "whites” (Macassans
and Europeans) in myth and oral history and the presentation of these representations by non-Aboriginal academics, and the way
they inter-relate.

See Rudder (1993:23) for an overview of the use of “inside” and “outside” terminolgy.

194

Who are the Bayini?

journey of the spirit of the Yolngu deceased
(Mountford 1956-64: 314) or through associa¬
tion with sacred totemic emblems, as in Figure 1.

The next wave of visitors was known as the
Bayini. The term was described by the Bemdts
(1954: 34),

(It) is used to classify the first (settlers on)
the northern coast of Arnhem Land...All along
the coast...special sites are said to have been
associated with them. There are certain rocks
that symbolise a wrecked Bayini boat, or a lost
anchor, and various places are named after
incidents that occurred during their stay. Per¬
sonal names of many Aborigines today signify
their Bayini derivation...We cannot be certain
who these Bayini people were...The Aborigines
are quite decided that (they) were not like the
Macassans who came after them because they
are remembered particularly for the golden
colour of their skin.

Unlike “white” Macassans, the Bayini are
said to have built stone houses in north-east
Arnhem Land, while the women made pottery
and cultivated the ground, growing rice in Gumatj
and Warramiri territory. Bemdt and Bemdt
(1949: 220) and Worsley (1955) give contrast¬
ing views on the involvement of local Aborigi¬
nal people in these industries.

Little information has been published on this
group and what there is has been gleaned from

Aboriginal song cycles (e.g. Bemdt and Bemdt
1947a: 135). Of the researchers with first hand
experience in north-east Arnhem Land, a majority
either doubted the historical existence of pre-
Macassans or used a number of terms to describe
their mythological character, e.g. Wangarr or
“Spirit”Macassans(Wamer 1969; Williams 1986).

While Bemdt (1965: 5) acknowledged the
difficulties of locating the Bayini in time and
space because of the way in which the Aborigi¬
nal past is enshrined in contemporary song and
ritual, he did not question informants' state¬
ments that a separate wave of contact was in¬
volved. He thought that the Bayini might have
been sea nomads from the islands off southern
Sulawesi, Selayar orBouton. The word Bayini is
in common use in those parts (Bemdt 1965: 5),
meaning a girl or woman. The Bemdts re¬
mained undecided, however, as to whether set¬
tlement was planned or enforced, but suggest
that contact could have been before the sixteenth
century, although they add that more archaeo¬
logical or other evidence is needed (Bemdt and
Bemdt 1947a: 133).

The Bemdts suggest Aborigines did not de¬
sire to imitate the Bayini, preferring their own
way of life. While the two groups co-existed,
they did not seem to have been willing to leam
from each other. The Bayini, for instance, kept
the secret of weaving to themselves (Bemdt and
Bemdt 1954: 38).

F.- 2 2 A Macassan prau under sail. Artist unknown, c. 1970. This “outside" or public painting depicts the annual historical
vovaees of Macassans to and from Arnhem Land. The black triangle represents the north wind Lungurrma . which brought the
praus on to the coast, and identifies the painting as belonging to the yirritja moiety or half of Yolngu society. Four Macassans
(at the oars and below deck) and an Aboriginal (on top) are on board. Photo courtesy of P. Lane.

195

I. McIntosh

Fig. 3. Macassan sailing canoe. Artist - Liwukang of the Warramiri clan. 1993. Trepang in the sailing canoe and on the sides
of the painting links it with the Macassan era, and not the Bayini, according to the artist. It is an “outside” or public painting
referring to historical encounters, and yet the octopus, a sacred totem of the artist’s clan, entwining the mast, hints at the “inside”
significance of these same encounters in Aboriginal law. Courtesy of the Museum and Art Gallery of the Northern Territory
(Abartl 175).

was purely commercial, for they themselves had

no use whatsoever for trepang.

The established picture presented in the lit¬
erature is that the Macassans and Aborigines co¬
existed, on the whole, in peace and harmony,
although it is difficult to substantiate such gen¬
eralisations (Macknight 1972:289; Swain 1993;
Worsley 1955: 9). According to Macknight
(1972: 303) there was a degree of mutual trust
between the groups, and certain Macassan boat
captains had a basic knowledge of Aboriginal
culture, languages and place names. It appears
that the visitors had no interest in changing Abo¬
riginal lifestyles.or of exploring inland rivers for
the purposes of colonisation (Warner 1969:449).

Aborigines assisted in the industry by diving
for trepang, smoking and curing the sea slug,
fishing, building smoke houses for curing, cut¬
ting firewood and digging wells (Worsley 1955:
3). Not only were they wage labourers, they also
stockpiled turtle shell, pearls and trepang and
exchanged these for trade goods such as axes,
knifes and tobacco (Macknight 1972: 308;
Worsley 1955: 3). As with the Bayini and “to¬
tem” hunters, images of specific Macassans are
rare. They appear as silhouette figures aboard
Macassan praus, as in Figures 2 and 3.

Representations of the Bayini in art are rare
and there is a degree of confusion between them,
Macassans and Aborigines. Inone case, aBayini
man identified by the Bemdts (1954: 36) as
Wonatjay is described by Mountford as an Abo¬
rigine that had been to Macassar (1956-64:
299). In another example, the image of Bayini
women at work (on their weaving looms in a
painting by Liwukang of the Warramiri clan in
Cawte 1993: 83) is identical in many respects to
aGumatj painting by Mungurrawuy made in the
1950s, though he describes the female figures as
Macassan.

After the Bayini, there were evidently two
phases of Macassan contact. The Bemdts (1954:
40) speculate that the first lasted from 1700 until
the 1820s and the second until 1907. Macknight
(1972: 289) however found little evidence to
justify this view. In giving an overview of the
Macassan trepang industry, Macknight (1976:
1) says:

It began in about AD. 1700 and continued on
until the early years of this century. For most of
the nineteenth century, and probably the hundred
years before that as well, at least a thousand men
made the voyage each year...their product was
exported to an international market. Their object

196

Who are the Bayini?

Little evidence is available from the early
period of Macassan contact, but the prodigious
wealth of the visitors and their desire to share
this with Aborigines (Worsley 1955: 3) and
accounts of rituals held jointly by the visitors
and Aborigines (e.g. Fig. 4) suggest mutual co¬
operation between the parties at some point in
the past. Bemdt and Bemdt (1954: 46) noted:

...(T)he most colourful of all...(Macassan)
ceremonies, so Aborigines say, was the gala
Bau’wulji, translated graphically by Mission
natives as “Christmas Day”. It was held during
the “cold” season at Manangu (Mununu or
Dholtji) in country called by the Macassans
Wusing-djaladjari, (Ujung Djalatjirri - Cape
Wilberforce) “the Last Point”- their last main

Fig. 4. Ritual of celebration. This “outside” painting by Mattjuwi of the Gumatj clan in 1993 depicts a scene from the early
Macassan era. Two Macassan praus are shown, and on the top left are items of trade. On the top right, Macassan men (with crosses
on their chests) and Aborigines, are seen performing a ceremony together. Courtesy of the Museum and Art Gallery of the
Northern Territory (Abart 1177).

197

I. McIntosh

Fig. 5. The Abduction. As with Figure 4, this 1993 painting
shows articles of trade and also a trcpang cooking area. On
board the boat is a lone person. Lul'warriwuy of the Ngaymil
clan, an Aboriginal woman abducted by the Macassan Captain
Maliwa from Arnhem Bay in the 1890s. She never returned.
She was a classificatory mother of the Gumatj artist. Mattjuwi
(Cooke 1986:33-34; Sydney Morning Herald , “Good Week¬
end". 2 October, 1993; 11-16). Courtesy of the Museum and
Art Gallery of the Northern Territory (Abart 1178).

Indonesians and their Aboriginal employees
joined in dancing and singing.

Curiously, the area where this ritual is said to
have occurred is in the very same place that
Matthew Flinders met the Macassan Pobasso in
1803. Pobasso reported little contact with the
Aborigines, believing them to be treacherous
(Flinders 1814). Bemdt and Bemdt (1954: 46)
locate this period of joint celebration in the early
stage of Macassan contact, and such a view lends
support to informants’ statements that there was
considerable difference in the nature of contacts
between the two phases. Thus while the
Macassans appear to have been welcomed ini¬
tially, relations deteriorated in the last phase of
contact. The Bemdts (1954: 47) commented:

In the old days...trading partnerships were
established between the two groups. These
involved reciprocal obligations and created
classificatory ties of kinship.

In the final stages of the industry, “...the
Aborigines began to feel that they were being
exploited" (Bemdt and Bemdt 1954: 110). Al¬
cohol was introduced as a means of paying
Aborigines, leading to violence on both sides. It
led to prostitution in the Macassan camps and
the death of many Aborigines by gun or sword
(Bemdt and Bemdt 1954: 47). The Bemdts add
that there were many murders of crew members
as well. The atrocities described by Searcy (1909,
1912) are linked to this time, and are reflected in
Figures 5 and 6.

By the late nineteenth century there was a
growing concern by Australian authorities about
the trepang trade. The Macassans were not
paying taxes and there were questions about
their treatment of Aborigines. In 1907, after
continued pressure for Europeans to take control
of the industry, the Government prohibited the
entry of these peoples and a major chapter in
Australian history came to an end (Macknight
1986: 73).

meeting place on the coast, prior to the home¬
ward trip with the south-east winds. This was
during the first phase of contact, before they
expanded their operations further into the west
of Arnhem Land. Here in the sheltered waters
behind Cape Wilberforce all the praus assem¬
bled, their cargoes were checked, and they held
a great ceremony of farewelL.they played mu¬
sical instruments, let off fireworks, and both the

BAYINI: PRE-MACASSANS, WANGARR
MACASSANS, OR SPIRIT MACASSANS?

Figures 1 to 7 contain images of what Abo¬
riginal people perceive to be actual historical
events and/or periods, e.g. the Bayini coming to
Port Bradshaw; joint ceremonies of celebration
with Macassans; trepanging; voyages to
Macassar; the abduction of women and so on. In

198

Who are the Bayini?

Fig.6.The Firing Squad.This 1993 pain tingby the artist Daypurryun of the LiagawumiiTclan also relates
specifically toan historical incident fromhis clan territory at Elcho Island. At thebottom one can see three
men (Aborigines) and a boy (a Macassan). They face a Macassan firing squad for the murder of a crew
of trepangers, but on the strength of evidence from the young man, the Aborigines are spared, and one clan
leader is taken to prison in Macassar (Isaacs 1979:84; Read and Read 1991:16-18; Sydney Morning
Herald, "Good Weekend”, 2 October, 1993: 11-16). Courtesy of the Museum and Art Gallery of the
Northern Territory (Abartl 176).

199

I. McIntosh

Fig. 7. The Bayini atPort Bradshaw. 1950s. Artist unknown. In one ofonly a few depictions of the Bayini, this“inside" or sacred
Gumatj/Warramiri painting depicts two Bayini men holding swords in eitherhand.and two Bayini women. The figures arebright
yellow in colour, which Bemdt and Bcmdt (1947a: 133) say is the same as that of a certain species of flying fox. Courtesy of the
Art Gallery of New South Wales.

this section, I look at the confusion that has
resulted from attempting to view Aboriginal
accounts of the Bayini or the Macassan past as
history alone.

The earliest anthropological studies in north¬
east Arnhem Land which mention outsiders
were undertaken by the American Lloyd Warner
in the 1920s and Donald Thomson in the 1930s,
and neither mentions the Bayini. Warner (1969)
wrote in detail of the social effects of Malay
(Macassan) contact and discussed the impact of
what he termed the Wangarr Macassans on
Aboriginal mythology, and in particular, mortu¬
ary ritu al. Thomson ’ s (1957) approach was some¬
what different. He raised questions of the psy¬
chological impact of early Macassans on Abo¬
rigines, but apart from a few lines on accultura¬
tion and the apparent hero worship of the early
traders, he did not pursue the matter. Thomson
(1949) looked instead at the impact of a “virile”

culture from Indonesia on local Arnhem Land
trade networks and speculated on the ancient
nature of the visits. As with Warner, there is a
presumption that Macassans were the only group
of visitors to have been involved with Aborigi¬
nes, and that complex mythologies were in place
explaining their origin and purpose.

It was in the 1940s that the term Bayini first
appeared in the literature, but it is evident that in
the various accounts they are seen not only as
Wangarr Macassans. but also as a wave of
historical influence prior to Macassans. Berndt
and Bemdt for instance say that the Bayini came
at the beginning of the historical period, after
the “golden age” of the creator beings.
Djang’kawu had already peopled the region but
the arrival of the Bayini was considered so early
that “...they are regarded less as historical fig¬
ures than as mythological spirits" (Berndt and
Berndt 1954: 33).

200

Who are the Bayini?

As Wangarr or “spirit” Macassans, the Bayini
are seen to be of a similar order to other totemic
or ancestral figures, i.e. creating sacred sites and
transforming the landscape, with Aborigines
acquiring theirqualities, characteristics and laws.
For instance Williams (1986: 2) says:

...when the Macassans first arrived on the
coast', the Aborigines then living there were
expecting them. This was because during that
time in the far distant past when spirit-beings
were investing the world with meaning, spirit-
Macassans had appeared. They brought with
them in spirit form the things that “real”
Macassans would later bring and explained
their use.

Mountford’s (1956-64:334) account includes
one of the very few myths published on the
Bayini. He tells of how golden skinned people
from Jumaina (Fig. 7) brought their women and
children with them to Arnhem Land. At the
Bayini totemic centre at Port Bradshaw there are
two totemic wells in which their ancestors still
live today. He writes:

Gurumuluna, the headman of die Bayini and
his sister camped at a waterhole...(at) Port
Bradshaw. After a while, transforming them¬
selves into flying foxes, the brother and sister
flew...(to the western shore) where they changed
themselves back into human beings and copu¬
lated. Both their footprints and the marks of the
woman's buttocks can be seen on the rocks at
Dalmumnia. Later, as flying foxes, they flew to
Melville Bay, where again, transforming them¬
selves into human beings, they lived as man and
wife (Mountford 1956-64: 336).

In the 1940s, Aborigines stressed the differ¬
ences between Wangarr Macassans (i .c. Bayini)
andMacassan trepangers(Bemdt 1965:5). Apart
from differences in terminology for common
items of trade and their different skin colour to
Macassans, the Bayini are said to have brought
women with them. No women were present on
the Macassan boats (Berndt and Bemdt 1954:
36), although one reference suggests that Asian
women might have been “given” to Aboriginal
men in exchange for access to land (Bemdt and
Bemdt 1947b: 249), but this was questioned by
informants I spoke with. Some stories, however,
suggest a relationship between Bayini and
Macassans. Bemdt and Bemdt (1954: 37-38)
for instance record that:

Before the Bayini left the Australian main¬
land, they looked around and saw smoke rising
from a fire far away at Macassar. Then the
Bayini headman spoke, “We have to go there
and leave this place. It is better that native
people should work for us there”. When they
reached Macassar the Bayini built a big tank
and filled it with hot water. They climbed into
it and soaped themselves until they became
white in colouring, just like (Macassans and)
Europeans. Aborigines say that this is why the
Bayini songs are mixed with those relating to
Macassans.

While R.M. Berndt (pers. comm. 21 August,
1989), saw the historical aspect of the Bayini
stories as being of primary significance, there
are other pieces of information which he re¬
corded which hint more at the material’s deeper
significance to the yirritja moiety and clan iden¬
tity. For instance, he notes that apart from the
yirritja secular ceremonies associated with the
Bayini , the Warramiri clan have sacred Bayini
ceremonies called Gwolwunbulma, Lilgarun,
Mara’raguma or Janderalguma, relating to the
“shovel-nose” iron-bladed spear, the knife and
axe, and in these, Bayini figures are used as a
madayin or rangga (sacred objects). Bemdt and
Berndt (1949: 221) also note that:

The Bayini ...sent out their ideas and cus¬
toms, (mainly ritual and ceremony), called
“law”, to different yirritja groups of aborigines,
from Cape Wilberforce to Caledon Bay.

Fig. 8. A line drawing of the “man of iron”, Birrinydji, from
Cawte (1993:78).

201

I. McIntosh

In the late 1960s historians set about debunking
what they deemed to be inventions in Aboriginal
accounts of the past. Macknight (1976), in his
work The Voyage to Marege, refers to the puz¬
zling ceremonies with Macassan associations:

Most remarkable of all are certain stories,
associated with particular places in Australia,
that I believe to be derived from experiences
and observation in South Celebes and possibly
elsewhere. The idea of things which properly
belong overseas has been transferred to famil¬
iar places in order to integrate this knowledge
into the spatially oriented framework of Abo¬
riginal thought" (Macknight 1976: 161).

He adds that the stories of the Bayini are, “...a
most remarkable instance of the need to distin¬
guish between the account of the past current in
a society and the actual events of the past,”
(Macknight 1976: 161), and that:

Whatever one makes of the Baiini stories,
there are no details in this material or in any
other Aboriginal information about the
Macassans that can be assigned with confi¬
dence to a period other than the eighteenth and
nineteenth centuries. The time needed to pro¬
duce the effects of Macassan contact seen in
some Aboriginal societies is a matter of opin¬
ion. I believe that two centuries is more than
enough time (Macknight 1976: 97-98).

The overwhelming view therefore is that the
stories of the Bayini are largely invention and
based on actual Macassan and perhaps other
contacts, and also Aboriginal experiences over¬
seas, but what is the Bayini law, and of what
relevance was it in the past or now? Why did
Aborigines insist in the separation of the Bayini
from Macassans, and why did they focus on the
activities of women in accounts of pre-
Macassans? In the next section I compare and
contrast two myths, one from the 1920s and the
other from the 1990s, as a means of examining
these issues.

COMPARISON OF THE EARLIEST AND
MOST RECENT REPRESENTATIONS IN
MYTH

The recent appearance in the literature of the
“man of iron”, Birrinydji (Fig. 8), (Cawte 1993:
42; McIntosh 1992: 101; 1994a: 78-79; 1994b:
93) has necessitated a complete re-thinking of

the ways in which the Aboriginal past has been
represented by academics. Birrinydji is both an
ancestor for the Warramiri clan, and the hus¬
band of a female ancestral being called Bayini,
and is the perceived source of the “other’s”
power. Birrinydji brought not only the historical
pre-Macassans but also Macassans and Europe¬
ans to Arnhem Land. Pre-Macassans, who also
go by the name Bayini, are linked with the
emergence of this “inside” or sacred law.

Take the following for example. Burrumarra (in
McIntosh 1992: 101) of the Warramiri clan says:

Birrinydji and Bayini are for Dholtji (a
Warramiri homeland). All things come from
Birrinydji. Two thousand years ago people came
to our land. They had a job to do. They wanted
to make the land and the people strong. It was
at Birrinydji's command that they came. The
iron in the ground acted like a magnet, drawing
them in.

Birrinydji was like a blanket over the land.
Everything came under him. He was both white
and black. He was very rich and had many
things. He was an iron-maker. Bayini made
clothing, planted rice and directed Yolngu
women in this. Birrinydji came from the ground,
from the gold beneath.

Visitors to Dholtji had settlements all along
Cape Wilberforce. There were thousands of
people, men, women and children...When we
followed Birrinydji's law, we prospered. But
then things started to go wrong. We wanted only
good but bad came too... We turned our back on
the laws of Birrinydji and we lost everything.
There is great sadness in our memory and this
is why we don't like to bring it up. Today we
follow the laws of Birrinydji. We have only the
song and the ceremony but we have lost the
ability to make iron. But if we follow this law,
maybe these tilings will come to us again.

Then compare this with a myth recorded by
Warner in the 1920s, which is the only other
reference to the “fall” in the literature. It is an
“outside” myth concerning the activities of the
Warramiri totemic DogBoTlili, and its meeting
with Macassans at the very same place referred
to above, i.e. Dholtji on Cape Wilberforce (D.
Burrumarra pers. comm. 1990). The informant,
not identified by Warner (1958: 537) told that:

A very, very long time ago everything was
different. People who lived in this place had
skin just like Macassar men and Macassar men

202

Who are the Bayini?

had skin like black men. Macassar men worked
for black men then, just like we work for
Macassar men by and by.

Dog was talking to his master. “We better
break this house down and throw him away and
live without houses."

The master of the dog talked to him. He said,
“What do you want? Do you want something?
He imitated the master’s speech.

The man said, “No, I asked you.”

Dog said, "No, I asked you.”

They repeated this several times. That dog
did not understand what that black fellow-
white man saying. The black man who was a
white man said,” You don’t understand what I
am asking you for.” He said that to the dog.

The dog continued acting silly, and said he
did not want anything. The white man said,
“You’re the black man now and I the white, lam
the master. I’ll give you matches and tobacco
and a sailing boat and tomahawks.”

The dog said, “1 don’t want them. You can
have them.”

That master went back and another headman
came. The first headman said, "I have come
back.”

The other said, “Why have you come back?”
He said, “ Because Dog talked badly to me.

I offered to give him all of those things and he
said he did not want them.”

The other Macassan man said, “All right, all
we people will keep all these things because Dog
talked that way. We’ll let them work for us.”

The Macassar men came in their boats for
trepang. The black people who belonged to Dog
went out to work for them. They became more
and more black because Dog had acted so silly.

A long time ago we people were white, now we
are black. The name of this time a long time ago
was Wangarr time. This happened when the
world started.

Given the lack of any historical evidence for
any other significant contacts, the implication is
that an “inside” view of Bayini secs the visitors
as an ahistorical population instigating the law
of Birrinydji at the “beginning of time”. The
Bayini era is both a time in which Aborigines
possess the wealth of the “other” and are in the
image of that “other”, and simultaneously, the
“other” ( Wangarr Macassans) are present on
Aboriginal land and their wealth, which Abo¬
rigines desire, is denied them. In a “cargo" type
perspective of “loss”, Macassans and Europeans

enjoy the wealth that Aborigines once possessed.
It also explains how one came to be in a subor¬
dinate position in relation to the other.

The “fall” is therefore crucial in understand¬
ing the distinction between the “historical”Bay/n/
(Wangarr Macassans) and Macassans. It is re¬
ally a distinction between the “inside” law of
Birrinydji and Bayini, and an associated sacred
past, and “outside” Macassan history, that pe¬
riod of contact with trepangers.

This is highlighted by the fact that in all
myths which mention Macassans, they are re¬
jected. Mountford (1956-64: 282) refers to a
bloody battle between the Thunderman and
Macassans, in which the latter .are turned away,
and Bemdt and Bemdt (1954: 89) tell of a
similar conflict over territory between the saw¬
fish and the visitors. In examining this rejection,
McIntosh (1992,1994a) looked at the somewhat
ambiguous Dog/Macassan “encounters” and
came to the conclusion that the two totemic
operators (Dog and Macassan) are seen to be of
the same cosmic class, i.e. being lawless or out¬
side the law. Bayini or Wangarr Macassan sto¬
ries, on the otherhand, represent sacred “inside”
Aboriginal law, and there is no equivalent bat¬
tles over entry or rejection by totemic beings.

So it is evident that the earliest and the most
recent accounts of “inside” law complement
each other. They both paint a picture of Aborigi¬
nal “loss” relative to the "other”. Burrumarra
says that neither he nor his brothers spoke
publicly of Birrinydji in the past. Explanations
were always given in terms of the historical
Bayini, and then only “women’s” stories were
revealed. It was because “all things from women
are free.” Birrinydji stories on the other hand
were “too strange, too strong, and too hard to
understand”.

Birrinydji is too deep in us, too close to our
heart. He is the King and we are his subjects.

We can talk of Bayini only. These Yolngu
women had many children who grew up to be
bunggawa. We are the children of those women;
the children of Birrinydji and Bayini. Bayini is
a Yolngu woman, Gumatj, Warramiri, Wangurri
and so on. Birrinydji is our father (D.
Burrumarra, pers. comm. 1990).

Belief in Birrinydji is for Aborigines alone,
Burrumarra says. It is not about Macassans or
Europeans, but rather who Aborigines are or
should be in relation to them, i.e. respected as
people of equal status if not wealth. In contrast

203

I. McIntosh

to these “inside” stories, “outside” representa¬
tions of contact have taken numerous forms and
in the next section I put forward a possible
explanation for the fact that accounts appear to
be in constant flux.

INTERPRETATIONS AND REVELATIONS
IN TRANSITION

Building on Worsley’s (1955: 9) view that
descriptions of the Macassan era are coloured by
the contemporary social situation and relation¬
ships with “whites”, my hypothesis is that
Birrinydji/Bayini mythology is constituted in
the ever-changing relations between Aborigines
and non-Aborigines, and its emergence and
decline in relevance correlates with develop¬
ments in this area.

In the 1920s, the recorded view was that spirit
Macassans had been present at the “beginning of
time”, and that Aborigines once possessed the
wealth that now only the “other” enjoys. In the
1940s and 1950s, these Wangarr beings were
referred to as the Bayini and were also seen as
actual historical voyagers. In the 1960s,
Macknight (1972: 312-313) found that knowl¬
edge of the Bayini had all but disappeared, with
detailed knowledge limited to a few older men,
in particular Burrumarra. In the 1990s, the
“inside” law of the past was openly discussed for
the first time, suggesting that the stories of the
Bayini, rather than being lost, had been ob¬
liquely referred to by Warner in the 1920s,
partially revealed to the Bemdts and Mountford
in the 1940s, and totally restricted in the 1960s.

Aborigines were not being deceptive in speak¬
ing only in terms of myth or history with anthro¬
pologists or historians, but rather were being
cautious about revealing “inside” details of a
body of law which has its foundation in the
knowledge of the existence of the “other” and
the presence of that “other” on Aboriginal land.

I have the advantage here of hav ing had as my
chief informant David Burrumarra 3 , a man who
worked with both the Bemdts, Thomson, and
Macknight, and who was the primary owner, as
leader of the Warramiri clan, of the stories
relating to the exploits of pre-Macassans, and
Birrinydji. In discussions with him, it is appar¬
ent that the distinction between “inside” and
“outside" myth/history has allowed for a diver¬

sity of ways of both speaking about Macassans
and also for making adjustments in what is to be
publicly known of the past.

Examples of this are numerous. Following
contact with academic historians, scientists and
missionaries, certain “outside” changes were
made in accounts of the trcpangers. Tamarind
trees contain the spirit of Birrinydji in “inside”
interpretations, but are also “outside” historical
markers of Macassan campsites. Similarly, an
“inside” view sees the dug-out canoe as the
legacy of Birrinydji, via contact with the histori¬
cal pre-Macassan Bayini, whereas in “outside”
accounts, they were often the gift of Macassan
trepangers at the end of the season (D.
Burrumarra, pers. comm. 1992).

Macknight (pers. comm. 1994) says he was
told various snippets of myth but was not in a
position to do anything with such data, and one
might presume that Aboriginal informants would
pick up on this very quickly. If the “Macassan”
past was to be investigated, then “totem” hunters
and the Bayini, or other sensitive matters, need
not be brought up in discussions.

The considerable changes in accounts in the
literature from the 1920s up to the present can
also be seen in terms of this management of “in¬
side” and “outside” interpretations of law by
Yolngu leaders. What can be spoken about de¬
pends to a large degree on the nature of relations
between the parties involved. Thus, when Warner
and Thomson were doing their fieldwork, the
Methodist missions were just beginning. Accord¬
ing to Burrumarra, the movement of people to
the communities ofMilingimbi (1923), Yirrkala
(1935) and Galiwin’ku (1942) was seen by some
as part of Birrinydji’s plan (McIntosh 1994b:
102). It was to be the fulfilment of Birrinydji’s
“promise”. Aborigines and Europeans would
share in the wealth of the land, and this might
explain the emphasis on the mythological in the
early accounts. Only later, in attempts to protect
“inside” knowledge from outside scrutiny, was
there motivation to create an “outside” histori¬
cal term for an “inside” body of law.

If one accepts the view that the necessity for
maintaining traditions relating to the perceived
motives and power of the “other” came into
question in the mission period, it is not therefore
surprising that there would be a corresponding
emphasis on “history” as opposed to myth when
“black” and “white” lived together in a single

3 David Burrumarra died on October 13, 1994, at Elcho Island.

204

Who are the Bayini?

Fig. 9. The Warramiri Flag Treaty Proposal. Burrumarra (centre) is flanked by Liwukang (left) and Berripang. Birrinydji is
shown on the bottom left, and on the right are two octopus emblems, suggesting links to the “outside” image (Fig. 4).

community. Just as followers of “cargo cults” in
Melanesia now deny they were ever involved in
such movements (Hermann 1992: 66), there is/
was also a degree of embarrassment associated
with older beliefs associated with Macassans,
i.e. that Aborigines were once “white”. Rather
than risk their being held to mockery, Birrinydji
and Bayini largely disappeared from general
usage. How else can one explain why research¬
ers in the 1960s up until the 1990s were not
shown or told about the major Bayini and
Birrinydji sites, which in many cases coincide
with trepanging areas?

Birrinydji ceremonies are still performed to¬
day at initiations, funerals, community celebra¬
tions and even in some cases, Christian gather¬
ings, but it is not a celebration of the past and
performance makes no direct reference to the
visits of Macassans. Along with sites in the
landscape and sacred objects connected to this
law, such rituals remain of fundamental signifi¬
cance to the identity of many Yolngu clan groups.

This is highlighted in Yothu Yindi’s 4 songs
“Treaty” and “ Djapana ”, which both refer to the
Birrinydji theme and to Yolngu solidarity in the
face of the non-Aboriginal presence (Burrumarra,
pers. comm. 1992).

BIRRINYDJI AND A TREATY PROPOSAL
IN NORTH-EAST ARNHEM LAND

So why has new data on Birrinydji and Bayini
been released in recent times? Certain events, I
suggest, have ensured Birrinydji’s continuing
relevance, even if detailed knowledge of this law
is restricted to a single generation of older men.
The mining operation at Gove, and the per¬
ceived flagrant denial or ignorance of Aborigi¬
nal rights by fishermen and tourists has seen the
re-emergence in new forms of action based
around the Birrinydji theme - in this case, a
treaty proposal for Australia by Burrumarra and
two other senior men of the Warramiri clan,
Liwukang and Wulanybuma (Figs 9-10).

4 A Yolngu “rock” band from north-east Arnhem Land.

205

I. McIntosh

The Warramiri plan was to make a series of
flags incorporating symbols from the past and
the present which would unite “black” and
“white” Australians. Burrumarra’s idea was that
in the future, the country would have not just
one, but many national flags, each containing
Aboriginal symbols relevant to the area in which
it was flying, depending on what it meant to the
Aboriginal inhabitants (Burrumarra, n.d.). For
instance if one was in Sydney, the flag might
incorporate images of the possum, rainbow ser¬
pent and dolphin or other designs. Various
versions of the Warramiri flag, for instance,
incorporate sacred images of the whale, squid,
octopus and Birrinydji. The common thing with
all the Australian flags, Burrumarra said, would
be that the “sacred" symbol of Great Britain, the
Union Jack, would appear in the upper left hand
corner, as in the current flag 5 .

In Burrumarra’s vision, the past and present
seen through Birrinydji!Bayini mythology cre¬
ates an image of a future in which Aborigines
will be recognised as spokespersons for the
country and have the riches of non-Aborigines.
What had happened on Aboriginal land al¬
lowed all Australians to reflect on what was
happening now in Aboriginal dealings with
non-Aborigines. From the earliest memories of
Warramiri history when “black” and “white”
people danced together, to the “fall”, to fights
against domination by outsiders, both Macassans
and miners, the Warramiri have come to an
understanding about how events should and
must proceed. As Burrumarra says, “the policy
we have in relation to the Balanda (non-Atx>
rigine) is the same now as it was in the begin¬
ning”. This is the policy of Birrinydji. It was
only by telling the story of the past that a
deeper understanding of the rights of Aborigi¬
nes can be understood and respected. The myths
are therefore not merely about a remote
Dreamtime or the result of reflections upon the
Macassan period. Birrinydji is about the future
and what actions need to be taken to make it
coincide with a utopian vision of how things
were prior to the “fall”.

The present living conditions of Aborigines
in Arnhem Land and the complex relationships
with non-Aborigines have warranted a change

design.

in the status of Birrinydji, heralding the need for
new laws, to be enacted by Parliament, to con¬
firm one another’s rights in relation to each
other. As Burrumarra says, if his treaty proposal
is accepted, and the Government raises the flag
representing both black and white Australians,
then “ Birrinydji can put down the swords”:

...we would still use them for ceremonies,
but we are burying them in the sand. We do not
live by the sword any longer, by that law. We
have different principles.

Today, people live as one group. “Black” can
marry “white” and vice versa. This is part of the
lesson of the treaty. We are different today than
before. We live by a new law. Our histories
have merged. The law of the past was Yolngu
for Yolngu and Bayini for Bayini. This is
Birrinydji's law. We do not mix. Outsiders
tried to steal the women and steal the land. We
would lose everything. But we can share the
future if there is equality.

We know the law of Birrinydji in the past but
this is a new world now... In the past we were
his servants, servants of the bunggawa
(Birrinydji), Now we want equality. We ask
Bill Hayden, Can we be equal in your eyes?
(Burramurra pers. comm. 1990).

The Warramiri leaders are thus putting into
effect “...in transformed economic and social
space, a Habitus which is the product of a
previous state in this world,” to use Bourdieu’s
(1993) terms. To re-use, in novel ways, beliefs

5 Here again we have two separate interpretations of the significance of a particular symbol. While there is a push to have the
Union Jack removed from the Australian flag in Australia, for the Warramiri it is a symbol of profound importance, and of
relevance in depicting and commenting on historical relationships between Aborigines and non-Aborigines.

206

Who are the Bayini?

all but “lost in time”, is an attempt at transform¬
ing the nature of practices associated with the
belief. In airing the stories, the elders are at¬
tempting to facilitate the enactment of a treaty of
reconciliation with Government authorities.

CONCLUSION

While interpretations of the Aboriginal past
make reference to varied contacts between Abo¬
rigines and non-Aborigines, a majority of the
stories associated with that body of “inside” law
now referred to as Birrinydji are drawn from
experiences and reflections of Macassan con¬
tact. The search for one group by the “outside”
term for this body of law, i.e. Bayini is therefore
misplaced. It is of far less significance than to
ask the question, why have Aborigines repre¬
sented the past as they have done.

What is the significance of these representa¬
tions now in Aboriginal dealings with the
“other”? The subject is an extremely complex
one. At present, the term Bayini is the name of
a female ancestral being of the Warramiri and
otherclans in north-east Arnhem Land, and also
a personal name for a female Aboriginal, mean¬
ing “white woman” (Zorc 1986). It can also be
used in reference topre-Macassans, although, as
stated, this is an “inside” view of limited cur¬
rency today. It was used in the past as a means of
not speaking about Birrinydji, the Wangarr
Macassan, and it allowed for a distinction, in
“time” between “inside” “timeless” laws and
accounts of Macassan trepangers. I use the word
“time” here in inverted commas, for Birrinydji
and Bayini are ancestors of Warramiri clan
members, not in a biological sense, but in a
deeply held and paradoxical view that in the
past. Aborigines and the “visitors” were united
in the laws of Birrinydji, but the failure of
Yolngu to follow this law has become an expla¬
nation forperceived inequality or“loss” in terms
of the “other”. So the people Aborigines dealt
with on a day to day basis in the Macassan
trepanging era necessarily came after those that
Aborigines were separated from at the “begin¬
ning of time”.

Such findings highlight the problem of mak¬
ing definitive statements about the Aboriginal
past. Truth is a matter of context. Birrinydji
mythology is constituted in the ever-changing
nature of relations with the “other” and provides
a reference point for representations relating to

the Macassan era. In paintings one can see the
full range of views, from actual historical epi¬
sodes but which depict aspects of Birrinydji's
technology (knives and trepang boilers), to the
predominantly mythological, i.e. the Bayini at
Port Bradshaw. At different times, depending
on the nature of relationships between Aborigi¬
nes and the “other”, “inside” myth as opposed to
“outside” history may predominate in public
discussion, while at others, “history” is the
primary means of referring to a sacred past.

The body of “inside” law symbolised by the
use of the expression Bayini, remains of pro¬
found significance to the Yolngu, being associ¬
ated with particular tracts of land and sacred
ceremonies. Wh ile it i s questionable as to whether
people still believe that Birrinydji is the founda¬
tion of the wealth of the “other”, such laws
continue to motivate action designed to affirm
the place of the Yolngu in relation to the “other”,
as it must have done in the past, though under
quite different circumstances.

ACKNOWLEDGMENTS

This paper was first presented in the Anthro¬
pology seminar series at the Northern Territory
University in August 1994. The author wishes to

thankD.Meams,C.C.Macknight,P.McConvell,

and an anonymous reader for their helpful com¬
ments in the re-working of the original draft,
and also to P. Lane, J. Cawte, and the Aboriginal
artists Liwukang, Daypurryun and Mattjuwi, for
the use of their artwork.

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McIntosh, I.S. 1995. Maluku “totem” hunters and
Sama-Bajau in north Australian Aboriginal my¬
thology. Australian Folklore 10:50-60.

Morphy, H. 1990. Myth, totemism and the creation of
clans. Oceania 60(4): 312-328.

Mountford, C.P. 1956-64. Records of the American
Australian scientific expedition to Arnhem Land.
Vol 1. Art, myth, and symbolism. Melbourne
University Press: Melbourne.

Read, P. and Read, J. 1991. Long time, olden time.
Aboriginal accounts of Northern Territory his¬
tory. Institute for Aboriginal Development Pub¬
lications: Alice Springs.

Rudder, J. 1993. Yolngu cosmology. An Unchanging
cosmos incorporating a rapidly changing world?
Unpublished PhD thesis. The Australian Na¬
tional University.

Searcy, A. 1909. In Australian tropics. George
Robinson and Co: London.

Searcy, A. 1912. By flood and field. Adventures
ashore and afloat in north Australia. G. Bell and
Sons, Ltd: London.

Swain, T. 1993. A place for strangers. Towards a
history of Australian Aboriginal being. Cam¬
bridge University Press: Cambridge.

Thomson, D.F. 1949. Economic structure and the
ceremonial exchange cycle in Arnhem Land.
Macmillan: Melbourne.

Thomson, D.F. 1957. Early Macassar visitors to
Arnhem Land and their influence on its people.
Walkabout 23(7): 29-31.

Warner, W.L. 1958/69. A black civilization. A social
study of an Australian tribe. Harper & Roe:
Chicago.

Worsley, P.M. 1955. Early Asian contacts with Aus¬
tralia. Past and Present 7: 1-11.

Williams, N. 1986. The Yolngu and their land.
Aboriginal Studies Press: Canberra.

Zorc, R.D. 1986. Yolngu-matha dictionary. School
of Australian Linguistics, Batchelor Northern
Territory.

Accepted: 10 January, 1995

208

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1995 12:209-214

Book Reviews

Sue Ebury, Weary: the life of Sir

Edward Dunlop, Viking, Ringwood, 1994,

ISBN 0 670 847607, viii + 709 pp, $45.00.

The outlines of Sir Edward Dunlop’s life are
probably familiar to many Australians. ‘Weary’,
as he was usually known, was a national hero.
Bom in Victoria in 1907, he became a brilliant
surgeon and an international rugby union player.
He served during the Second World War as an
Army medical officer in Palestine,Greece, Crete,
Tobruk and Egypt before the Japanese captured
him in Java in 1942. His care for men working on
the infamous Burma to Thailand ‘death railway’
made him a living legend. After the war he was
a pioneer in the treatment of cancer and cared for
former prisoners of war. He was also involved in
various other community activities. At the time
of his death in 1993 he much honoured and held
in high regard.

There is, though, another version of Dunlop.
At least some of his medical colleagues resented
what they claimed was the high-handed way in
which he undertook his surgical work. Nor was
he always held in great respect among former
prisoners of war. Some of the latter, including
my father and several of his friends, criticised
him for self promotion, arguing that he was just
one of many medical officers who did useful
work in prison camps.

Sue Ebury is clearly among Dunlop’s most
ardent admirers. Her huge biography was writ¬
ten with Dunlop’s full cooperation. The pace is
often breathless and the prose is gushing. Ebury,
the New Zealand bom wife of a British peer, has
no doubt as to her subject’s greatness. She ac¬
knowledges some deficiencies in his character
but argues that these were of minor consequence
i n comparison with h is vast ach ievements. Dun lop
emerges from her pages as a source of inspira¬
tion. Not surprisingly, the prison camp years are
dealt with in great detail. Dunlop’s skills in or¬
ganising medical services under the most appall¬
ing conditions and his constant defiance of his
captors quite rightly get special prominence. One
can only be impressed at the story of devotion to
duty, courage and passion that Ebury relates.

The problem with Weary is that its exhaustive
and uncritical attention to Dunlop’s doings re¬
sults in the portrait of a man with some pretty
unpleasant characteristics and marked flaws of
character. In his youth he was an unashamed
militarist. While a university student he was
involved in physical attacks on other students
who did not share his own right wing political
views and supported hideous initiation ceremo¬
nies at his residential college. Though obviously
expert in his chosen profession, he sometimes as
a young medical practitioner obviously felt that
a boisterous social life was equally important.
During the Second World War, both before and
after his capture, he took needless risks that
placed his own life, and sometimes the lives of
others, in jeopardy. Married at the end ofthe war,
he very frequently neglected his wife and family
in the years that followed: he spent as much time
away from home as he possibly could. He was
notorious for his unpunctuality in surgical ap¬
pointments and his lack of consideration for
medical colleagues. He drove his car fast and
dangerously. His favourite sports, rugby union
and boxing, attracted him due to their ‘physical’
aspects.

The comments above may appear carping and
uncharitable but there is a great deal of detailed
evidence in Ebury’s book to support them. Dunlop
was, of course, a product of the era in which he
grew up and a very conservative profession.
Many readers of Weary will no doubt support
Ebury’s admiration of this ‘man’s man’. Others,
though, may share my disquiet.

David Carment

Department of History,

Northern Territory University,

Darwin, NT, 0909, Australia.

Dr. David Carment is Associate Professor of
History at the Northern Territory University and
has published widely in the areas of Northern
Territory politics, mining and cultural heritage.
His most recent publication is the History and
Landscape in Central Australia and he is soon to
publish a complementary volume about Darwin.

209

Book Reviews

Ross Fitzgerald, “Red Ted”: the life of
E.G. Theodore, University of Queensland
Press, St Lucia, 1994, ISBN 07022 2649 1,
xxiv + 487 pp, $24.95.

This is an outstanding biography of an ex¬
traordinary Australian. Its author. Associate Pro-
fessorRossFitzgeraldofGriffith University, has
written highly acclaimed texts on various as¬
pects of Australian history and politics. He has a
special interest in Queensland’s past and is also
well known as a commentator on political and
social issues. Edward GranvilleTheodore, known
widely as ‘RedTed’, was born in South Australia
in 1884. Largely self-educated, he became active
as a young man in the Queensland labour move¬
ment and within ten years was an outstanding
Queensland Premier. He later moved into fed¬
eral politics, becoming Treasurer in the Scullin
Labor government in 1929. A royal commission
into his mining interests forced his temporary
resignation and he lost his seat in the 1931
elections but he impressed many observers with
his considerable economic skills during the very
difficult Depression period. He subsequently
helped launch the Consolidated Press empire,
owned gold mines in Fiji and became a leading
administrator during the Second World War,
dying in Sydney in 1950. Fitzgerald’s account of
his life is meticulously researched, beautifully
crafted and full of insights into the character of a
man who was widely misunderstood.

There is so much in “RedTed” of interest that
it is impossible in a short review to do it justice.
One of its outstanding features for me is the
attention it devotes to Theodore’s very impor¬
tant north Australian connections. At Arltunga
near Alice Springs in 1903 he operated a hand¬
held drill and successfully convinced other men
in this very isolated goldfield to demand a pay
increase. In 1906 in the Chillagoe-Irvinebank
area near Caims he also mixed mining with
organisation of workers. With others in 1907 he
formed the Amalgamated Workers’ Association
of North Queensland, which later merged with
the powerful Australian Workers’ Union. In 1909
he was elected to represent the Chillagoe area in
the Queensland Legislative Assembly. Despite
his rapid rise to Premier in 1919, he remained
very interested in north Australian matters. He
wrote many articles on his proposed scheme for
a ‘Seventh State’ that would encompass the
Northern Territory and its surrounding areas.

His enthusiasm for the north received a rebuff in
1925 when he was defeated as a candidate for the
north Queensland federal electorate of Herbert
and the ‘Mungana affair’which did him so much
political damage in 1930 concerned his financial
interests in a north Queensland mining venture.
Fitzgerald argues convincingly that many alle¬
gations made about Theodore’s involvement with
Mungana were unsound and malicious yet con¬
cedes that ‘the taint of Mungana stayed with
Theodore until his death in 1950, and well be¬
yond’. As Director of the Allied Works Council
between 1942 and 1944, he organised resources
for vital war-related projects in north Australia.

Another fascinating aspect ofFitzgerald’s work
is the manner in which it sympathetically yet not
uncritically deals with Theodore’s personal life.
His paternal grandfather, Georges Teodorescu,
was a Romanian Orthodox priest yet Theodore
himself became a nominal Catholic and ulti¬
mately an atheist. Theodore’s marriage was, for
the most part, unhappy. His wife was often
difficult and demanding. Although actually
eleven years older than Theodore, she claimed to
be only two years his senior. Ultimately they
separated and Theodore had a relationship with
a woman in Fiji. He was not a man who often
revealed his emotions.

For most of his career Theodore was a moder¬
ate and a pragmatist who would not be out of
place in the Australian Labor Party of the nine¬
teen nineties. Buthe was also, Fitzgerald stresses,
‘much beyond the average run of Australian
politicians’. It is a tragedy that he was struck
down as a political leader at a time when his
particular skills were so desperately needed.

David Carment

Harry Gordon, Voyage from shame:
the Cowra breakout and afterwards,
University of Queensland Press, St Lucia,
1994, ISBN 0 7022 2628 9, xi + 331 pp,
$16.95.

It still comes as a shock to many Australians
when they are told that the largest prison escape
in history occurred near Cowra in New South
Wales on 5 August 1944. More than eleven
hundred Japanese prisoners of war tried to break
out of their camp. While twohundred and thirty-
five, including four Australians, died, hundreds

210

Book Reviews

escaped and were recaptured soon afterwards. As
Harry Gordon explains, the ‘full facts’ of the
escape were not disclosed until quite recently.
The author of an earlier book on the incident
published in 1978, he found in preparing a new
work that there was much information of which
he had not been previously aware.

Gordon is a'distinguished journalist who is the
author of nine previous books, most of which
have been well received. It is, then, not surpris¬
ing that Voyage from Shame is clearly written
and well organised. 1 found it difficult to put
down as the story it recounts is full of drama and
human interest. The book is also based on thor¬
ough research. Much relevant archival material
was used in its preparation and many of those
involved with the escape, both Australian and
Japanese, were interviewed. Gordon makes a
sincere effort to understand the Japanese partici¬
pants and hopes that his book will assist the
process of reconciliation. My one complaint is
that some dialogue, while based on evidence, has
clearly been invented for dramatic effect.

A point that emerges frequently is the enor¬
mous shame associated with surrender for the
Japanese. Many prisoners at Cowra had given
false names when captured so that dishonour
would not be brought to their families. The
breakout was, from an Australian perspective, a
futile business as there was virtually no chance
that any escapee could elude recapture for very
long. For the Japanese involved, though, the aim
was to remove sliame by being killed or commit¬
ting suicide. Even today some survivors of the
breakout find it difficult, if not impossible, to
discuss or even acknowledge their prisoner of
war experience.

Among the more prominent organisers were
airforce men captured in the Northern Territory.
Hajime Toyoshima, for instance, made an emer¬
gency landing on Melville Island after the Japa¬
nese raid on Darwin on 19 February 1942. He
was found by Matthias Ngapiatilawai, a young
Tiwi man who became the first person to arrest
a Japanese serviceman on Australian soil. An¬
other group of five airmen, including Marekuni
Takahara, were also washed ashore on Melville
Island after being shot down at sea nearby.

That the escape took place at all was due to lax
Australian procedures. Many members of the
garrison battalion at the Cowra camp were unfit
for normal military service and their weapons
were obsolete. There were some early warnings
that a breakout might take place but these were

not properly understood. Two middle-aged Aus¬
tralian soldiers died heroically in a vain attempt
to stop the escapees with a single Lewis Gun.
Even the efforts to recapture prisoners were
sometimes badly organised. Some soldiers sent
out to look for the Japanese were only armed
with bayonets.

Voyage from Shame deserves a wide reader-
ship. It ought to appeal to all those interested in
the Australian experience in the Second World
War. It also is worth close attention at a time
when many Australians are looking closely at
developing understanding with various Asian
peoples. Of these the Japanese are perhaps the
most important yet remain the most perplexing.

David Carment

Denis Winter, 25 April 1915: the inevi¬
table tragedy. University of Queensland
Press, St Lucia, 1994, ISBN 0 7022 2472 3,
xiv + 290 pp, $18.95.

Denis Winter is an historian with a special
interest in the First World War whose recent
book Haig’s Command: A Reassessment attracted
much criticism and led several reviewers to ques¬
tion his capacities as a researcher and writer. 25
April 1915 is slightly less controversial but some
of its claims are still likely to be contested. The
purpose of his latest work is to present a new
picture of the landing at Gallipoli, an event with
almost spiritual significance in Australia. Winter
claims to have used documents never before
studied by historians. He argues that these con¬
clusively demonstrate that the campaign was
doomed from the very beginning. This view is,
of course, in opposition to the quite widely held
belief that the Gallipoli operation may have
succeeded if the men in charge at various levels
had made wiser decisions during the campaign.

Winter considers various events leading up to
25 April through discussion of the soldiers in¬
volved, strategic planning and operational mat¬
ters. He argues that there was ‘a quantity of
forethought and planning in stark contrast to the
customary presentation of Gallipoli as a knee-
jerk response to the Russian appeal for assist¬
ance of January 1915’. He provides an evocative
description of the day to day experiences of the
Australians and New Zealanders training at Mena
in Egypt, not far from the great pyramids, and

211

Book Reviews

later at the island of Lemnos. There is extensive
use of their diaries and letters. As in Bill
Gammage’s classic The Broken Years , one gets
the impression of young men who were danger¬
ously naive. They often saw the war as a great
adventure with little realisation of the horrors
that would occur. Meanwhile, the politicians and
senior commanders were, Winter contends, ‘mud¬
dled’ in their thinking about the Gallipoli ven¬
ture and ‘mercilessly pushed forward’ with in¬
herently faulty schemes.

The most interesting part of the book deals
with the landing on 25 April. Winter suggests
that the true story is difficult to recount due to the
secrecy enforced by military commanders and
the fact that so many vital documents are miss¬
ing. He argues, though, that three distinct and
consecutive schemes existed for the landing,
each with a clear tactical logic behind it. The last
of these, contrary to the usual version in studies
of Gallipoli, indicated that Anzac Cove was the
chosen landing point and the troops did not
arrive there by mistake. Winter also contests
prevailing views about the soldiers’ conduct at
Gallipoli. Some formations disintegrated under
Turkish firepower, with men straggling back
towards the beach. Other soldiers, of course,
fought heroically. A special bond was formed
among those who survived the fighting on the
first day at Gallipoli.

Winter writes well and his book is easy to
follow. There are some excellent photographs
and maps. The book also appears to be thor¬
oughly researched although there are occasional
factual errors. I am not sure that he adds as
greatly to the Gallipoli story as he claims or
whetherexpert military historians will think much
of some of his arguments. For the general reader
interested in Australians’ experience of war,
though, 25 April 1915 is certain to be of interest.

David Carment

Gail Reekie, ed. On the edge: women’s
experiences of Queensland. University
of Queensland Press, St Lucia, 1994. ISBN
0702224618 RRP $29.95.

The title of this edited collection of essays. On
the Edge , relates not only to the marginalised
position of women, but of all Queenslanders in
relation to Victoria and New South Wales.

Queenslanders, it would seem, are searching for
new identities through their histories and litera¬
ture. Work began on this book in 1989 and it is
reasonable to speculate that after a year of cel¬
ebrating the bicentennial of the settlement of
Sydney, which became synonymous with the
settlement of all Australia, the collaborators were
keen to ensure other voices were heard in the
construction of our national identity.

The bibliographic survey concludes that
women have been excluded so far from most
general histories of Queensland and this text
therefore can be read as a contribution to the
search for a revised and feminised Queensland
identity. The masculinist images of the frontier
days no longer reflect the way Queenslanders
view themselves. Several of the writers identify
David Malouf, Jessica Anderson and Thea Astley
as the most influential figures in reconstructing
Queenslanders’fictional identities. Also signifi¬
cant, argue many of the writers, is the distinctive
architecture of the Queenslander house and the
way various spaces were created and used by the
occupants, and by women in particular.

There are thirteen essays covering sources,
histories, spaces, and politics and parties. The
majority of the contributors, of whom all are
women, were connected in some way with Griffith
University at the time of publication and the
camera-ready copy was produced with the as¬
sistance of the Division ofHumanities at Griffith
University, which makes this book a truly col¬
laborative effort. The col lection is presented as a
contribution to women’s studies rather than to
women’s history, though most of the essays use
historical methodology and the book is pub¬
lished under the University of Queensland’s
Studies in Australian History series.

The Introduction by Gail Reekie, addresses
the relevance of region to feminist enquiry. Re¬
gion, argues Reekie, is defined by borders con¬
structed for the use of geographers, demogra¬
phers and politicians.

Because women were primarily located in the
domestic and private sphere as daughters, wives
and mothers and only temporarily or informally
(if ever) as entrepreneurs or paid workers, the
majority were isolated from the effects of such
public and formal regional boundaries. Is region,
then, a usefu 1 concept in the analysis of women’s
relationship to history, culture and government?
(P-2.)

Reekie suggests that for regionalism to be
relevant to feminist study, the borders must be

212

Book Reviews

redefined. She describes this collection as the
beginning of a dialogue about the connections
between women, space and time. My one criti¬
cism of this collection, is that in raising these
issues in the Introduction, the reader expects the
essays to explicitly address these concerns but
most often this is not the case.

There is a tendency in most of the essays to
assume women’s oppression and to celebrate
women’s resilience, often without evidence.
There is, for example, only one reference to the
possibility of genuine partnerships between men
and women in pioneering endeavour. I found
Aline Gillespie’s essay, which describes how the
advent of the Young Women’s Christian Asso¬
ciation (YWCA) into the Lockyer Valley en¬
riched the lives of young women in the region
between 1935-40, one of the most powerful
because of her skilful use of evidence. Gillespie
discusses the limited options forcompanionship
and leisure for so many of the girls who were
unable to continue at school beyond grade eight
because their labour was needed to support their
families. Those living “off the bitumen” were
isolated further by unreliable black soil roads
and lack of transport. In the time before teenag¬
ers were invented, these young women were
neither children nor married women and appar¬
ently in an hiatus. Gillespie’s use of oral evi¬
dence to describe the marginalisation of these
young women, is compelling. The YWCA pro¬
vided a variety of Saturday afternoon entertain¬
ments including picnics, tennis, craft activities
and most importantly opportunities for compan¬
ionship.

Readers in the Northern Territory will recog¬
nise many of the themes in On The Edge, and will
no doubt grin wryly at the notion of
Queenslanders regarding themselves as
marginalised. More importantly, however, this
collection suggests that women’s experiences
are unrestrained by traditional regional borders,
and that landscape, space and time are more
likely to define women’s domains.

Julie T. Wells

Department of Education,

Northern Territory University,

Darwin, NT, 0909, Australia.

Julie T. Wells is currently working as
Reasearch Assistant to the Dean of Education,
NTU and is about to complete a PhD on the
assimilation policy in post-war Darwin.

Patricia Grimshaw, Marilyn Lake,
Ann McGarth, Marian Quartly,
Creating a nation, McPhee Gribble, South
Yarra, 1994, ISBN 0869140957, 360pp,
$16.96

It is sometimes said that both men and women
created this nation and that it is foolish to argue
otherwise. Men, it could be said, provided much
of the brawn and, fora century or more, the brain,
in the paid workforce; women saw to it that there
was another generation to inherit and carry on
their efforts. By and large, this has been the
conventional view of Australian society and
Creating a Nation, authored by four women who
are senior academics, Pat Grimshaw, Marilyn
Lake, Ann McGrath and Marian Quartly, seeks
to challenge that assumption. The authors note in
the introduction: ‘this books starts from the
premise that gender is integral to the processes
that comprise the history of Australia - that politi¬
cal and economic as well as social and cultural
history are constituted in gendered terms.’ As a
product of collaboration it is impressive; indi¬
vidual authorial style remains yet the book itself
is wholly integrated. Creating a Nation is valu¬
able in identifying not only the changes which
have taken place to women’s condition since
1788 but the effect on many of the women who
endured them. There are timely lessons to be
learnt from it.

The book begins, appropriately, by locating
the history within the context of Aboriginal
experiences of British colonisation. It is a meas¬
ure of the pervasive nature of the masculine
Anglo colonial construction of Australian his¬
tory that the reader is continually disconcerted
by the new interpretations offered by these aca¬
demics; at timesbewildered, appalled orcharmed
with the slightly guilty feeling of female focus.
This is a big book on a big subject, which
philosophically and methodologically is taking
on a powerful construction established in two
hundred years of political and social history. It is
difficult to do Creating a Nation justice in a
small review but I hope that I have make it clear
that this is a significant and important work.

I found most interesting and heartening those
sections dealing with twentieth century events
where women’s voices become stronger and
more confident, as they, usually in times of
national peril, were permitted a more active role
within society. The dichotomy between the roles

213

Book Reviews

of working woman and mother is well illustrated;
for the most part neither role a matter of personal
choice. The drudgery of unpaid or menially paid
work and multiple children was the lot of all
women, regardless of education or background.
Today’s woman needs to knowhow hard fought
was the battle to become an entity in her own right
and not simply a chattel of a male, be he father or
husband; she also needs to know how difficult
was the road (andhow many women were cruelly
treated in the male-dominated system along the
way) to easier divorce and support from the state
in times of crisis.

The title of the book does not do it justice;
nations are created by lines drawn on maps.
Children are created, nations are constructed;
but it is the creation of children which cause
many women so much joy and so much pain. The
construction of Australian society as predomi¬
nantly Anglo-male, for women too, leads to pain
but little joy. Women are used to effacing them¬
selves and being effaced from public profile
except under rigidly circumscribed construc¬
tions. Here in the NorthemTerritory, in a remote
capital, we are used to reading national histories
where ourregion is ignored. It is a measure of our

maturity as a community that we now share the
confidence to celebrate diversity outside the
stereotypes of gender, geography or politics.
The nation has been created by women and men,
and only a knowledge of women’s role in Aus¬
tralian history, such as found in this book, will
give validity to the choices which Australian
women inevitably make. Similarly, this book
marks a smal I but significant step down the long
journey towards a balanced view of Australian
society where all voices, no matter how quietly
spoken, are heard.

Helen J. Wilson

do History Department

Museum and Art Gallery of the Northern

Territtory,

P.O. Box 4646, Darwin, NT 0801, Australia.

Helen J. Wilson is a freelance historian who
is currently engaged on a variety of projects and
assists atMAGNTwith research on the historical
photographic collection. She has just completed
a women’s history ofsites in the Wharf Precinct,
Darwin, for the Northern Territory Government’s
Women’s Advisory Council.

214

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CITATIONS AND REFERENCES

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Brake, B„ McNeish, J. and Simmons, D. 1979. Art
of the Pacific. Oxford University Press: Wel¬
lington.

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The

Records Of The Museums & Art
Galleries of the Northern Territory

December 1995

Contents

ERRATUM - Mislabelling of figures of holotypes in one of the Bruce papers in Vol. 11.ii

CLARK, L.G., WENDEL, J.F. AND CRAVEN, L.A. - A new species of Micraira (Poaceae:

Micraireae) from northern Western Australia...1

WILLAN, R.C. AND CATTANEO-VIETTI, R. - New data on Chelidoneura amoena Bergh,

1905 (Opisthobranchia: Cephalaspidea: Aglajidae).9

LARSON, H.K. - A review of the Australian endemic gobiid fish genus Chlamydogobius, with

description of five new species.19

WELLS, A. - Larva, pupa and notes on general biology of Tinodes radona Neboiss (Trichoptera:

Psychomyiidae)...53

BRUCE, A.J. - Latreutes anoplonyx Kemp, 1914 (Crustacea: Decapoda: Hippolytidae), a jelly¬
fish associate new to the Australian fauna.61

LANSBURY, I. - Notes on the genus Anisops Spinola (Hemiptera - Heteroptera: Notonectidae)

of the Northern Territory and Western Australia.65

HORNER, P. - Two new species of Ctenotus (Reptilia: Scincidae) from the Northern Territory.77

AKERMAN, K. AND BINDON, P. - Dentate and related biface points from northern Australia.91

BRUCE, A.J. AND COOMBES, K. - The palaemonoid shrimp fauna (Crustacea: Decapoda:

Caridea) from the Cobourg Peninsula, Northern Territory. 101

MURANO, M. - Two new species of the genus Anisomysis (Crustacea: Mysidacea) from northern

Australia. 145

BERGQUIST, P.R. AND KELLY-BORGES, M. - Systematics and biogeography of the genus

lanthella (Demospongiae: Verongida: Ianthellidae) in the south-west Pacific. 151

DEL CERRO, L. AND LLORIS, D. - A new species of Lepidotrigla (Scorpaeniformes:

Triglidae) from the waters off northern Australia. 177

IZAWA, K. - A new fish parasite (Copepoda: Siphonostomatoida: Caligidae) from the Timor

Sea, Australia. 185

MCINTOSH, I. - Who are the Bayinil . 193

BOOK REVIEWS:

CARMENT, D. - Weary: the life of Sir Edward Dunlop.209

- "Red Ted": the life of E.G. Theodore.210

< ] - Voyage from shame: the Cowra breakout and afterwards.210

- 25 April 1915: the inevitable tragedy.211

WELLS, J.T. - On the edge: women's experiences of Queensland.212

WILSON, H.J. - Creating a nation.213

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